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International Code of Nomenclature

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Bacteria and Viruses

BACTERIOLOGICAL CODE

Iowa State College Press, Ames, Iowa, U.S.A.

International Code of Nomenclature

of
Bacteria and Viruses

BACTERIOLOGICAL CODE

^\ Publication Date: June, 1958

Edited by

The Editorial Board of the

INTERNATIONAL COMMITTEE ON BACTERIOLOGICAL

NOMENCLATURE

Approved by

The Judicial Commission of the International Committee

on Bacteriological Nomenclature

The International Committee on Bacteriological Nomenclature

The International Association of Microbiological Societies

The Plenary Session of the VI International Congress of Microbiology,
Rome, Italy, September, 1953

© 1958 by the Judicial Commission, Inlernational Committee on Bacteriological
Nomenclature of the International Association of Microbiological Societies. All rights
reserved. Composed and printed by The Iowa State College Press, Ames, Iowa, U.S.A.

Library of Congress Catalog Card Number: 58-9480

Foreword

Microbiologists who have occasion to use the scientific names of the
microorganisms with which they deal generally prefer to use correct
names and to use them correctly. Relatively few authors have special
or direct interest in the problems of nomenclature as such, but there
is general recognition that acceptance of the same names by various
authors is essential in a field such as microbiology which has
probablv more economic implications than any other subdivision of
biolog\ . One is confronted with the fact that the names given to micro-
organisms have been proposed by individuals whose major interest has
been the organisms themselves, not their names. Their economic sig-
nificance has commonly been stressed. These minute organisms were
found in some cases to produce disease in man, animals or plants; their
study became basic to the professions of medicine and veterinary medi-
cine; other microorganisms produced fermentation, decay and spoilage;
it was found that fundamental studies of celhdar physiology and
metabolism, cell structine, inheritance, enzymology, photos) nthesis,
production of antibiotics, preservation of foods and feeds, public
health, sanitation, soil fertility, plant pathology, and other fields re-
quired some basic knowledge of bacteriology. Those who discovered
and worked with these organisms recognized the need of giving names
to them, but frequently had little or no experience in scientific
nomenclature. What rules should be followed in the coining of these
names? Precedents to be followed were not clearly formidated in the
early days of bacteriology.

Carl von Linne (Linnaeus) in the latter part of the eighteenth
century proposed certain nomenclatural principles which were adopted
with surprising unanimity by biologists of his day. Later the
botanists and zoologists in separate international meetings and con-
gresses developed two codes of nomenclature, which agieed in most
points but differed in some. Many bacteriologists followed the Botanical
Code, some the Zoological Code, and others named the organisms
which they discovered with scant attention to established rules. It

[v]

vi International Code of Nomenclature

Foreword

became evident that rules in Botany formulated primarily by those
interested in the taxonomy of flowering plants, ferns and mosses did
not fit too ^\cll the needs of the bacteriologist.

THE FIRST INTERNATIONAL MICROBIOLOGICAL CONGRESS (1930)

J'he desire that s]>ecial attention should Ix- paid to the
peculiar needs of bacteriology was voiced at the First International
Congress of Microbiology convened in Paris in 1980 by the Inter-
national Society for Microbiology under the auspices of the Pasteur
Institute. As the result of recommendations made by several of the
delegates to the Congress, a Commission on Nomenclature and
Taxonomy was constituted to prepare and report recommendations
to the Plenary Session of the Congress.

The members of this commission were E. Pribram, Chicago,
U.S.A., Chairman: A. R. Prevot, Paris, France, Secretary: R. E.
Buchanan, Ames, Iowa, U.S.A.; K. Kisskalt, Germany; J. C. G.
Ledingham, London, England; Reiner Miiller, Koln, Germany; R.
St. John-Brooks, London, England, and I. Yamasaki, Fukuoka, Ky-
ushu, Japan.

Several resolutions prepared by the Commission were approved
unanimously by the Plenary Session. These resolutions (in their
English text) were as follows:

/. The founding of the International Society for Microbiology and
the establishment of Congresses of Microbiology make possible
for the first time adecpiate international cooperation relative to
certain problems of microbial nomenclature. It is clearly recog-
nized that the living forms with which the microbiologists con-
cern themselves are in part plants, in part animals, and in part
primitive. It is further recognized that insofar as they may be
applicable and appropriate the nomenclatural codes agreed upon
by international Congresses of Botany and Zoology should be
followed in the naming of microorganisms. Bearing in mind
however the peculiarly independent course of development that
Bacteriology has taken in the past fifty years and elaboration of
special descriptive criteria which bacteriologists have of necessity
developed, it is the opinion of the International Society for Micro-
biology that the bacteria constitute a group for which special
arrangements are necessary. Therefore, the International Society
for Microbiology has decided to consider the subject of Bacterial
Nomenclature as part of its permanent programme.

//. The International Society for Microbiology is of the opinion that
the interests of bacterial nomenclature will best be served by
placing the subject in the hands of a single International Com-

Bacteria and Viruses

Foreword

mittee, under the aesjis of the International Society for Micro-
biology, adequately representative of all departments of Bac-
teriology, on which experts from all spheres of bacteriological
research may work together. It is recognized that the subject of
bacterial nomenclature is of so wide a nature that unless the
personnel of an International Committee formed to deal with it
is representative of all aspects of bacteriology, it is not likely to
carry weight. Such a representative committee, to be called the
Nomenclature Committee for the International Society for Micro-
biology, is hereby authorized and constituted.

///. The Nomenclature Conmiittee for the International Society for
Microbiology shall be constituted as follows:

a. Two permanent secretaries shall be elected: one primarily to

represent medical and veterinary bacteriology, the other pri-
marily to represent the other phases of bacteriology. The
following individuals are hereby appointed secretaries.

(1) To represent primarily medical and veterinary bacteri-
ology Dr. Ralph St. John-Brooks, Lister Institute. London,
England.

(2) To represent primarily the other phases of bacteriology
Dr. R. S. Breed. Geneva, New York, U.S.A.

Should a secretaryship become vacant, the position may be filled
pro tempore by choice of the Committee. A permanent secretary
should be chosen by action of the next succeeding International
Congress for Microbiology.

b. The remaining members of the Committee shall be appointed
by such National Committees of the International Society and
by such of the various National Societies affiliated with the
International Society as may desire representation thereon.
Not more than three members may be thus chosen to repre-
sent a single nation. In addition, in order that the Committee
shall be truly representative of all interests, the Committee
is authorized to add such members as may be deemed de-
sirable.

IV. The duties of the Nomenclature Committee shall include the
following:

a. Through the secretaries the members of the Committee shall
be circularized with reference to such jDroblems of bacterial
nomenclature as may arise, and shall endeavour to reach an
agreement. No action relating to nomenclature shall be con-
sidered complete and operative until it has been considered
by all members of the Committee, until adequate publicity
has been given with respect to actions proposed, until approval

v/ff International Code of Nomenclature

Foreword

has been given by a majority of two thirds of the members of
the Committee, and until a report has been made to the next
succeeding International Congress for Microbiology and op-
)3ortunity thereby given for objection, modification or re-
jection by action of the Congress,
b. The Committee shall consider, among others, problems such
as criteria to be employed in classification, adoption of names
for species and genera conservanda, type species (including
their identification and preservation) , the encouragement of
monographing of special groups or genera of bacteria by those
l^est qualified to do the work, the enlargement of the scope
and usefulness of the various type culture collections by more
adequate support, and the preparation and publication of
such Committee and Subcommittee reports as mav be advisable.

r'. Copies of these resolutions shall be submitted to the appropriate
sections of the International Botanical Congress, Cambridge,
1930. It is the hope of the International Congress for Micro-
biology that the members of the International Botanical Congress
who are interested in bacterial nomenclature will see the advis-
ability of the special questions of nomenclature of bacteria being
considered by a single international authority and that they will
suggest names of members of the Botanical Congress willing to
serve on the committee who, in their opinion, would add to its
strength and authority.

VI. In view of the adequate provision made for special regulations
relating to the bacteria, and the feasibility of designating genera
conservanda among the bacteria by international agreement, it is
believed that the greatest stability will be conferred by the adop-
tion of the publication of Species Plant nriim by Linnaeus in 1753
as the point of departure for bacterial nomenclature. The adop-
tion of this date is recommended. It is further suggested that no
present action be taken with reference to a list of genera con-
servanda for the bacteria.

HI. Among the most important agencies working toward satisfactory
nomenclature and classification of bacteria are the several type
culture collections. These constitute invaluable repositories and
much of the future development of bacteriology will depend upon
their adequate growth, support and utilization; in some cases at
least they should develop into research institutes of high grade.
It is urged that the coordination and cooperation existing among
these institutions be extended the better to serve the interests of
bacteriology in its theoretical, medical and other economic aspects.
It is further urged that all bacteriologists publishing descriptions
of new species or important strains of bacteria deposit pure cul-

Bacteria and Viruses ix

Foreword

tures of such with a culture collection that they may be made
available to others interested. Particularly is it urged that the
adequate financial support of these culture collections by official
agencies, by educational and research instittitions and by the re-
search foundations constitutes an important and immediate need.

It will be noted that in the action of the Congress the develop-
ment of an adequate Bacteriological Code was linked with the Botani-
cal Code. The specific suggestion was made that members of the
International Botanical Congress, 1930, be apprised of the resolu-
tions passed by the First Microbiological Congress and that the
Botanical Congress be asked to cooperate. This was done, and the
two secretaries of the International Nomenclature Committee for
Bacteriology (Dr. R. St. John-Brooks and Dr. R. S. Breed) were
designated by the Botanical Congress as a special committee on the
nomenclature of bacteria.

THE SECOND INTERNATIONAL CONGRESS FOR
MICROBIOLOGY (London, 1936)

The International Committee met during the sessions of the
second International Congress for Microbiology in London in 1986.
Proposals by R. E. Buchanan and H. J. Conn to conserve the generic
name Bacillus Cohn 1872, to designate as the type species Bacillus
subtilis Cohn 1872, and to fix the type or standard culture as the
'Marburg strain' were appro\ed by the Committee and by the Plenary
Session of the Congress.

A further specific action of the Nomenclature Committee and of
the London Congress had to do with the duplication of generic names
in the Protista, the group ordinarily defined to include the protozoa,
algae, fungi and bacteria. Inasmuch as bacteria are usually inchided
among the plants, and subsequent plant homonyms are regarded as
illegitimate, the principal interest is the suppression as illegitimate
later homonyms in the protozoa and the bacteria. Prof. F. Mesnil
proposed and the Nomenclature Committee and the Congress agreed
that generic homonyms are not permitted in the group Protista;
fiuther that it is advisable to avoid homonymy amongst Protista on
the one hand, plants or animals (Mefazoa) on the other.

The Committee and Congress also acted favorably on a proposal
by Prof. R. S. Breed relative to non-capitalization of specific epithets
in names of species of bacteria.

"Bacteriologists should accept Article 13 of the International

Rules of Zoological Nomenclature, as follows:

'While specific substantive names derived from names of per-
sons may be written with a capital initial letter, all other
specific names are to be written with a small initial letter.' "
At this 1936 (London) meeting of the International Committee it

X International Code of Nomenclature

Foreword

was agreed that, bcloie the convening of the third International
Congress of Microbiology to be held three years later in New York,
a tentative Code of Bacteriological Nomenclature should be drafted
and presented for the consideration of the Committee. To facilitate
easy conference an American (Canadian and U.S.A.) Subcommittee
was constituted to prepare such a tentative code. The members of
this Subcommittee were R. E. Buchanan, Chairman; Robert S.
Breed; J. Howard Brown; 1. C:. Hall; W. L. Holman; E. G. D.
Murray; and Otto Rahn.

The chairman was asked to assemble material for consideration
by the members. A mimeographed brochure of 119 pages was pre-
pared imder the title "Rules of Nomenclature, Annotated". It con-
sisted of two parallel columns. In the first column the International
Rules of Botanical Nomenclature, including Principles. Rules.
Reco7nmendations, Notes and Examples were printed. In the second
column were listed suggestions for a code of Bacteriological Nomen-
clature formulated by making such minor modification of the Botani-
cal Code as seemed desirable, as by dropping of inapplicable sections.
In numerous footnotes were given the pertinent sections of the In-
ternational Rules of Zoological Nomenclature and the American
Code of Entomological Nomenclature. This material was sent to all
members of the Subcommittee and to a large number of other bac-
teriologists, including members of the International Committee in-
sofar as they could be reached. Criticisms and suggestions were in-
vited. More than 30 sets of comments and suggestions were received.
These comments were broken up into sections corresponding to those
of the suggested code, and the proposed code and comments again
submitted to the members of the Subcommittee in the form of a
mimeographed booklet under the title "Suggestions and Comments
on 'Rules of Nomenclature, Annotated' ". A new series of connnents
and suggestions was secured from the numerous collaborators,
tabulated and submitted once more to the Subcommittee. A final
revision was prepared to present to the International Committee at
its New York meeting in 1939. The text of this tentative code dif-
fered from the basic Botanical Code principally in the following:

a. A reorganization of the text of the code under the following

headings.

1. General Considerations; 2. General Principles; 3. Rules of
Bacteriological Nomenclature with Recommendations; 4. Pro-
visions for interpretation and modification of rules.

b. Elimination of items and sections of the Botanical Code which
seemed inapplicable to bacteriology.

c. Simplification where possible through rephrasing.

d. Selection of examples where possible from bacteriology.

Bacteria and Viruses xi

Foreword

THE THIRD INTERNATIONAL MICROBIOLOGICAL CONGRESS
(New York, 1939)

The proposed tentative code was considered at some length by
the International Committee for Bacteriological Nomenclature at
its New York meeting; many suggestions developed. The report was
also presented to one of the sections of the Congress, and about one
hundred copies of the "Annotated" and "Tentative" codes dis-
tributed.

Upon recommendation of the International Committee on
Bacteriological Nomenclature the Plenary Session of the Third In-
ternational Congi-ess for Microbiology on Sept. 9, 1939 approved the
following resolution:

1. That a recognized Bacteriological Code be developed.

2. That publication of such a proposed Code when developed be
authorized with the proviso that it shall be regarded as wholly
tentative, but in the hope that it shall be widely tested so that
it may be brought up for further consideration and final dis-
position at the next Microbiological Congress which should
normally take place in 1942.

3. That the Nomenclature Committee, as at present constituted,
shall continue to function under the auspices of the Interna-
tional Association of Microbiologists* as it did under the Inter-
national Society for Microbiology.

4. That the International Committee shall select from its member-
ship a Judicial Commission consisting of twelve members, ex-
clusive of members ex officio, and shall designate a Chairman
from the membership of the Commission. The two Permanent
Secretaries of the International Committee on Bacteriological
Nomenclature shall be members ex officio of the Judicial Com-
mission. The commissioners shall serve in three classes of four
commissioners each for nine years, so that one class of four
commissioners shall retire at every International Congress. In
case of resignation or death of any commissioner, his place shall
be filled for the unexpired term by the International Committee
at its next meeting.

The functions of the International Committee on Bacteriological
Nomenclature were more accurately defined as follows:

a. To consider and pass upon all recommendations relating to
the formation or modification of Rules of Nomenclature. The
Committee will recommend such action as may be appropri-
ate to the next Plenary Session of an International Congiess

* The new name approved for the international organization sponsoring micro-
biological congresses.

xii Internafional Code of Nomenclature

Foreword

for Microbiology.

b. To consider all Opinions rendered by the Judicial Commis-
mission. Such Opinions become final if not rejected at the
meeting of the International Committee next following the
date on which the Opinion was issued.

c. To designate official Type Culture Collections.

d. To receive and act upon all reports and recommendations
received from the Judicial Commission or other committees
relating to problems of nomenclature or taxonomy.

e. To hold at least one meeting triennially in connection with
the meeting of the International Congress for Microbiology.

f. To report to the final Plenary Session of each Congress a re-
cord of its actions, and to recommend for approval such ac-
tions as require the approval of the Congress.

g. To cooperate with other Committees, particularly those of
the International Botanical and Zoological Congresses, to
consider common problems of nomenclature.

The functions of the Judicial Commission of the International
Committee on Bacteriological Nomenclature were also defined as
follows:

a. To issue formal Opinioyn when asked to interpret rules of
nomenclature in cases in which the application of a rule is
doubtful.

b. To prepare formal Opinions relative to the status of names
which have been proposed, placing such names when deemed
necessary in special lists, such as lists of Nomina Conservanda,
Nomina Rejicienda, etc.

c. To develop recommendations for emendations of the Inter-
national Rules for Bacteriological Nomenclature for pre-
sentation to the International Committee.

d. To prepare formal Opinions relative to types, particularly
types of species and genera, and to develop a list of bacterial
genera which have been proposed ^vith the type species of
each.

e. To prepare and publish lists of names of genera which have
been proposed for bacteria, for protozoa, or for other groups
in which microbiologists are interested in order to assist
authors of new names in avoiding illegitimate homonyms.

f. To develop a list of publications in microbiology whose
names of organisms shall have no standing in bacteriology in
determination of priority.

g. To edit and publish the International Rules of Bacteriologi-
cal Nomenclature, Opinions, Lists of Nomina Conservanda,
Nomina Rejicienda, Type Species, etc.

h. To report to the International Committee at its triennial
meetings all Recommendations, Transactions and Opinions.

Bacteria and Viruses xiii

Foreword

i. To report to the Imernational Committee at its triennial
meetings the names of all Commissioners whose terms of
service expire, likewise a list of all vacancies caused by resig-
nation or death.

Recommendation. Whenever, in the opinion of any micro-
biologist an interpretation of any rule or recommendation
is desirable because the correct application of such a rule or
recommendation is doubtful, or the stability of nomenclature
could be increased by the conservation or by the rejection of
some name which is a source of confusion or error, it is
recommended that he prepare a brief outlining the problem,
citing pertinent references and indicating reasons for and
against specific interpretations. This brief should be submit-
ted to the Chairman of the Judicial Commission; if desired,
through one of the Permanent Secretaries. An Opinion will
be formulated, which may not be issued until it has been
approved by at least eight members of the Commission.
It was further voted:

That the Proposed International Rules of Bacteriological
Nomenclature, in so far as they have been developed by the
American-Canadian Committee on Compilation of Proposals
on Bacteriological Nomenclature for the International Com-
mittee and modified by action of that Committee, shall be
referred for final emendation and publication to the Judicial
Commission in accordance with Provision (c) above as re-
corded.
The minutes of the International Committee contain the follow-
ing statements relative to the Judicial Commission:

With regard to the constitution of the Judicial Commission,
members of the Committee present were requested to ^i\e its
Secretaries lists of persons that they wished to nominate as
members of the Judicial Commission, and the Secretaries
were requested to transmit such nominations to the entire
Committee for ballot, giving members the option of sub-
stituting other names if they so desired. It was agreed that
after the final ballot the four persons receiving the greatest
number of votes should be elected for the nine-}ear period
and that the four persons receiving the smallest number of
votes should be elected for the three-year period. The re-
maining four are to serve for a six-year period.
Nominations to membership on the Judicial Commission were
made by the membership of the International Committee in attend-
ance at the New York meeting. The Permanent Secretaries then
conducted a mail ballot, resulting in the election of twelve members
(Commissioners) and designation of R. E. Buchanan as Chairman.

xiV International Code of Nomenclature

Foreword

R. S. Breed and R. St. John-Brooks as Permanent Secretaries of the
International Committee also became ex officio members and Perma-
nent Secretaries of the Commission.

The records of the Congress showed a membership of 62 on the
International Committee on Bacteriological Nomenclature as of
August 1939. There were representatives of Microbiological Societies
of 24 nations as follows: Argentina, Australia, Belgium, Brazil, Bul-
garia, Canada, Denmark, Deutsches Reich, Eire, France, Great Britain,
Holland, Hungary, Italy, Norway, Palestine, Poland, Roumania, Spain,
Sweden, Switzerland, United States of America, Union of Soa iet
Socialist Republics, and Uruguay.

It was expected that the mandate of the Congress to the Judicial
Commission to develop and publish a tentative proposal for a Code
of Bacteriological Nomenclature would be followed promptly. The
final determination of the constitution of the Judicial Commission
itself was long delayed because of the outbreak of World War II
while the New York Congress was in session. It soon proved im-
practicable to circulate copies of the nomenclatural proposals and
to secure comments from all members of the Commission.

Dr. Ralph St. John-Brooks of the Lister Institute, London, one
of the Permanent Secretaries of the International Committee in
March 1942, spent some days with the Chairman of the Commission
in conference and in editing the manuscript which had been re-
viewed by the Committee at the New York City meeting.

THE FOURTH INTERNATIONAL MICROBIOLOGICAL CONGRESS
(Copenhagen, 1947)

The Proposed Bacteriological Code of Nomenclature as author-
ized by the Third International Congress for Microbiology was
printed in June 1947 in a limited edition for distribution and for
use by the Judicial Commission and the International Committee at
the Fourth International Congress in September 1947.

At the Copenhagen Meeting the proposed Code was considered,
revised, and approved for publication by the Judicial Commission,
the International Committee and the Plenary Session of the Con-
gress. The English text was published in March 1948 in the Journal
of Bacteriology, later (Sept. 1949) in the Journal of General Micro-
biology. A Spanish translation (1949) by Prof. Verna was published in
Argentina in De Archives de Farmacia y Bioquimica del Tucuman and
a German translation by Dr. med. Hubert Bloch (1950) in the
Schweizerische Zeitschrift fiir allgemeine Pathologie und Bakteri-
ologie. A French translation by Dr. Pre\ot and a Japanese translation
were also issued.

Bacteria and Viruses xv

Foreword

THE FIFTH INTERNATIONAL MICROBIOLOGICAL CONGRESS
(Rio do Janeiro, 1950)

Meetings of the Judicial Commission and of the International
Committee were held at Rio de Janeiro and Petropolis (Brazil) in
August 1950. Among the important actions of these bodies, con-
firmed by the Plenary Session of the Congress, were the following:

1. An Editorial Board was established consisting of the Chairman
of the Judicial Commission and the two Permanent Secretaries.

2. Publication of a quarterly "International Bulletin of Bacterio-
logical Nomenclature and Taxonomy" was authorized; to be
edited by the Editorial Board.

3. Agreement was reached that some revision of the International
Bacteriological Code of Nomenclature was desirable and the
Judicial Commission instructed to incorporate certain amend-
ments approved, and to prepare recommendations for the 1953
International Microbiological Congress to be held in Rome.

The actions taken by the Commission, the Conmiittee and the
Fifth Congi'ess are reported in Volume One "The International
Bulletin of Bacteriological Nomenclature and Taxonomy."

In preparation for the Rome Congress (September 1953) the
provisional agenda for the meetings of the International Committee
and of the Jtidicial Commission were prepared and published in the
June (1953) issue of the International Bulletin.

THE SIXTH INTERNATIONAL MICROBIOLOGICAL CONGRESS
(Rome, 1953)

The Editorial Board prepared a series of "Proposals Relative to
Emendation and Publication" of a revised International Bacterio-
logical Code of Nomenclature (International Bulletin 1953 pp. 31-
63) which recommended publication of the revised Code, suggested
that the Rules and Recommendations be adequately annotated, and
that there be noted significant resemblances to the Botanical and
Zoological Codes of Nomenclature and likewise important differences
between them. The hope was expressed that texts in other languages
could be published simultaneously with the English text. In all, sixty
draft proposals for amendment, deletions and modifications of the
Code were submitted and acted upon.

The Judicial Commission, through the Editorial Board, was di-
rected to edit, annotate, and publish the Code as finally approved
by the International Committee and the Plenary Session.

The name of the Code was fixed as The International Code of
Nomenclatine of Bacteria and Viruses.

The manuscript for the Code in original draft form, including
Annotations and Appendices, was submitted for editorial suggestions

xvi International Code of Nomenclature

Foreword

to all members ot the Judicial Commission and to about twenty-five
bacteriologists experienced in nomenclature and taxonomy. Un-
fortunately the preparation of the text and annotations has been so
time-consuming that it has not been possible to include texts of the
Code in the several important languages of science. It is to be hoped
that this may be done in future printings.

REFERENCES

Buchanan, R. E. 1939 (February) (Editor) . Rules of Nomenclature: Annotated;
with Suggestions for Rules of Bacteriological Nomenclature. Prepared for
the American-Canadian Committee on Compilation of Proposals for Consider-
ation by the Third International Congress for Microbiology. Mimeographed pp.
118. Ames, Iowa. U.S.A.

1939 (July) (Editor) . Proposed International Rules of Bacteriological

Nomenclature. Prepared for Consideration by The American-Canadian Com-
mittee on Compilation of Proposals on Bacteriological Nomenclature for the
International Committee. Mimeographed, pp. 63. .Ames, Iowa. U.S.A.

, and Ralph St. John-Brooks. 1947 (June) (Editors) . Proposed Bacteriological

Code of Nomenclature. Developed from proposals approved by International
Committee on Bacteriological Nomenclature at the Meeting of the Third Inter-
national Congress for Microbiology. Publication authorized in Plenary Session,
pp. 61. Iowa State College Press Ames, Iowa. U.S.A.
, and Robert S. Breed 1948 (March) . (Editors) . International Bacter-

iological Code of Nomenclature. Journ. Bact. 55: 287-306. Also reprinted in
September 1949 Journ. General Microbiology 3:444-462.

Verna, Luis C. (Translator) 1949. Codigo International de Nomenclatura Bacteri-
ologica. Archivos de Farmacia y Bioquimica del Tucuman. ^: 283-3 16. Tucuman,
Argentina.

Bloch, Hubert (Translator) 1950. Internationaler bakteriologischer Nomenklatur-
codex. Schweiz. Zeitschr. allgem. Path. u. Bakteriologie. 13: 358-383. Basel,
Schweiz.

Editorial Board. 1953. The International Bacteriological Code of Nomenclature:
Proposals relative to emendation and publication. Internatl. Bull. Bact. Nomen-
clature and Taxonomy 3: 31-62. Iowa State College Press, Ames, Iowa. U.S.A.

Cowan, S. T. and T. Wiken (Secretaries) 1953. Minutes of the Judicial Commission
Meetings held at Rome in Connection with the VI International Congress for
Microbiology. September, 1953. Internatl. Bull. Bact. Nomenclature and Tax-
onomy 5:141-154. Iowa State College Press, Ames, Iowa. U.S.A.

1953. Minutes of Meetings of the International Committee on Bacteriological

Nomenclature held at Rome in Connection with the VI International Congress
for Microbiology September, 1953. Ibid. 5:155-161. Iowa State College Press, Ames,
Iowa. U.S.A.

Preface

The histor) ot the development of the 1958 Revised Edition of the
International Code of Xomenclatiue of Bacteria and Viruses has
been given in the Foreword. Here it is fitting that there be
acknowledgement of the generous assistance given by many individuals
and organizations in the preparation and editing of this Code.

The task of developing a wholly satisfactory Bacteriological Code
is not complete. New problems involving nomenclature of the bacteria
will arise and will require solutions. There have as yet been no final
recommendations and no conclusions as to what special Rules and
Recommendations will be needed to make functional any proposals
to be made by the International Subcommittee on Taxonomy of the
Viruses relative to virus nomenclature. The increasing use of term-
inologies applicable to strains and grotips of bacteria of infrasubspecific
rank makes necessary careful study of the best methods for preventing
confusion, even some degree of nomenclatural chaos, in the naming
of taxa of lower rank than subspecies. The growing recognition of
the value of the type concept in standardization of names may mean
the incorporation into the Code of a definition of Type Cidture Col-
lections and their functions in stabilization of bacteriological nomen-
clature.

A reading of the Annotations of the several Rules and Recom-
mendations of the Bacteriological Code reveals a variance in term-
inology (sometimes in basic concepts) in the three Biological Codes
of Nomenclature (Botanical, Zoological and Bacteriological) . These
differences have come about through the peculiarly independent de-
velopment and history of Botany and of Zoology. The organization
which can facilitate any attempt to reconcile these interdisciplinary
differences must represent biology as a whole and on an international
basis. The International Union of Biological Sciences would seem
to be the agency able in some effective manner to develop fruitful
consultations among the nomenclatural commissions of the three
disciplines.

The Editorial Board and the Judicial Commission are most grate-
ful for the generous subventions that have made possible publication
of this revised Bacteriological Code. Organizations particularly help-
ful have been the International Union of Biological Sciences, the

[ xvii ]

xviii International Code of Nomenclature

Preface

Society of American Bacteriologists, and the Society for General Micro-
biology. The Iowa State College has likewise been most generous in
its provision of office facilities.

The Editorial Board is grateful also for permission given h\ the
Commissions concerned to quote from the International Code of
Botanical Nomenclature and from the International Code of Zoological
Nomenclature where it has been desirable to compare resemblances
and differences between these Codes and the text of the revised Inter-
national Code of Nomenclature of Bacteria and Viruses. However, the
final text of the International Code of Zoological Nomenclature had
not been adopted in final form at the time of publication of the Inter-
national Code of Nomenclature of Bacteria and Viruses (June, 1958) .
In consequence some quotations may not represent final action by the
1958 Zoological Congress. If there are here included unintentional
misinterpretations, they will be corrected in later editions of the Bac-
teriological Code.

The manuscript for the Code in original draft form, including
annotations and appendices, was submitted for editorial suggestions
to all members of the Judicial Commission and to about thirty other
bacteriologists experienced in nomenclature and taxonomy. The
suggestions received were reviewed by the Judicial Commission. The
Code represents a high degree of international cooperation. The Edito-
rial Board wishes to express its real appreciation for the helpful co-
operation received.

TJie Editorial Board

R. E. Buchanan, Chairman S. T. Cowan, Secretary

T. WiKEN, Secretary W. A. Clark, Secretary

(resigned 1 April 1957) (appointed 8 October 1957)

Table of Contents

Foreword v

Preface ........... xvii

Chapter 1. General Considerations ^3

1. Scope of Bacteriological Code of Nomenclature. 3

2. Principles, Rules, and Recommendations. Definitions. 5

3. Provisions for establishment of an International Committee on
Bacteriological Nomenclature and of its Judicial Commission. 6

Chapter 2. Principles 9

1. Essential points in nomenclature. 9

2. Significance of estai:)lished custom in bacteriological nomenclature. 9

3. Interdependence in bacteriological, botanical, and zoological
nomenclature. 9

4. Names of all taxa to be Latin or Latinized. 11

5. Nomenclature deals ^vith categories of taxa, terms designating
names of taxa, and with the individual names of taxa. 12
Note 1. Definition and use of 'taxon' (plural "taxa") .

2. Definition of "name," of "legitimate" and "illegitimate"
as applied to names and epithets, of "correct name," of
"effectively published" and of "validly published."

6. Scope of Bacteriological Code. 13

7. Terms denoting rank of taxa: names of categories. 13

8. Primary purpose of naming taxa. 15

9. Principle of priority in determining correct names. 15

10. Principle of stability in nomenclature. 16

11. Significance of nomenclatural types. 17
Note. Definition of "nomenclatural type."

12. Principle of valid publication of names of taxa. 18

Chapter 3. Rules and Recommendations 21

Section 1. Names to be applied to taxa of difFerent ranks. 21

1. Names of suprageneric taxa, of classic origin or Latinized. 21

2. Names of taxa above the rank of family, how formed. 21
Rec. Preferably based on type genus of a contained family. 21

3. Names of taxa between suborder and genus. 22

4. Suffixes used to designate suprageneric taxa. 24

5. Names of genera and subgenera, how formed. 26
Rec. Suggestions to authors of new generic names. 30

[xix]

International Code of Nomenclature

Table of Contents

Name f)t a species;
specific epithets.

I)inarv com I)

How toiiiicd. KiiKJs ot

Note. Definition of "epithet."

Rer. a. Parenthetical position of name of subgenus in name of a
species.

b. Suggestions to authors of new specific epithets.

c. Use of genitive and adjectival forms of a specific epithet
(in the same genus) .

7. Name of a subspecies; a ternary conil)ination. How formed. Equiva-
lence of "subspecies" and "variety."

8. Infrasubspecific subdivisions. How designated.

Rec. Definitions of strain, type, group, phase, form, state, and stage.

Section 2. Designation of nomenclatural types.
9a. Each taxon has a nomenclatural type. Definition.
9b. Designation of a nomenclatural type for a suprageneric taxon.
9c. Designation of a nomenclatural type for a genus or subgenus.
9d. Designation of a nomenclatural type for a species or sidjspecies.
Rec. a. Desirability of designating nomenclatural types of newly

named taxa.
h. Preservation of

type" material.

Section 3. Publication of names.

10. Beginning date for valid publication of names of bacterial taxa.

11. Conditions of "effective" publication.

12a. Definition of "valid" and "validly published."

12b. Label on a cidture or preparation not valid publication.

12c. Names proposed provisionally or not accepted by author not
validly published.

12d. Listing a name as a synonym not valid publication.

12e. Listing of subordinated taxa not valid publication.

12f. Date of a name is that of its valid publication.

Rec. a. Diagnoses in unfamiliar languages.

b. Publication should be precisely dated.

c. Validity of publication in reprints.

d. Rank and location of new taxa should be indicated.

e. Nomenclatural types shoidd be designated.
/. Etymology of new names desirable.

13. Conditions for valid publication of generic and subgeneric names.

Note. Monotypic new genera.

Rec. Combined generic and specific description.

H. Conditions for valid publication of names of species and sub-
species.

Section 4. Citation of names of authors and taxa.

15. Citation of name of author and date of publication.

Rec. a. Designation of emendations.

b. Citation of description by one author in publication of
another.

33

Bacteria and Viruses

Table of Contents

16. Citation of author when rank of taxoii is changed. 76

Rec. a. Citation of name pul)lished as a synonym. 78

b. Citation ni synonymy of name invalidated by earlier
homonym. 78

c. Citation of nomen nudum. 78

d. Citation of a nonieti conservandum. 78

e. Spelling of names cited in s\nonymy. 78'

Section 5. Changes in names of taxa. 81

17a. Changes in names, when permissible'- 81

17b. Changes in name of a genus when divided. 81

17c. Changes necessitated by exclusion of type. 81

18. Changes in names of a species or of a subspecies when divided. 83

19. Rule of priority when taxa are united. 84
Rec. Choice of name when two generic names are of same date. 84

20. Name of subgenus which includes the type species of the genus. 85

21. Name of subspecies which includes the type of the species. 85

22. Retention of names of taxa with change in rank. 86
Section 6. Rejection and Replacement of Names. 89

23. Retention of inappropriate or disagreeable names of taxa or of
epithets. 89

24. Rejection of illegitimate names. Definition of illegitimacy. 91
Rec. Generic names that are later homonyms of generic names in
zoology. 93

25. Rejection of illegitimate specific and subspecific epithets. 95

26. Replacement of illegitimate names and epithets. 97

Section 7. Orthography and Gender of Names. 99

27. Retention of original spelling. Orthogiaphic variants. 99
Note 1. Definition of "original spelling."

2. Use of alternative or wrong connecting vowels in com-
pound words.

3. Reference to Judicial Commission.

•/. Acceptan.ce of author's correction of error.

5. Correction of spelling of Greek names incorrectly trans-
literated.

6. Connecting vowels in compound words.

7. Correction of spelling of a name or epithet should be
used with reserve.

8. Preservation of intentional Latinizations of names of
persons.

9. Suppression and transcription of diacritic signs.

10. Authorization for use of w, y, and k in Latinized words.

Rec. a. Correct transliteration of Greek to Latin. 105

/;. Choice of faulty or alternative transliterations. 105

c. Different font desirable for published scientific names. 105

d. Derivation of name of new genus or subgenus from the
name of a person. 107

e. Derivation of specific or subspecific epithet from name of

a man. 109

/. Derivation of specific or subspecific epithets from name of

women. 1 1 1

xxii International Code of Nomenclature

Table of Contents

g. Retention of original spelling and correct Latinization. 112

h. Specific and non-specific epithets are not to be capitalized. 112

/. Connecting vowels in compound names and epithets. 113

). Etymology of new generic names and epithets should be

given. 115

k. Choice between alternati\e spellings of names and epithets. 115

/. Varying transliterations from languages not using Latin

alphabet. 116

m. Correction of spelling of names and epithets having dia-
critical marks. 116

n. Use of names and epithets without Latin endings (non-
declinable words). 116

o. Transliteration of Greek names. 116

p. Abbreviation of generic name. 117

2S. Gender of generic names, how determined. 118

Chapter 4. Provisions for modification and amendment of
Rules, and for an International Committee on Bacterio-
logical Nomenclature .123

1. Procedure for Modification of Rules. 123

2. Lists of genera conseruanda. 123

3. Lists of genera rejicienda. 124

4. Permanent International Committee on Bacteriological Nomen-
clature. How constituted. Membership. Recognition of alternates.
Functions of International Committee. Election and duties of Chair-
man and Vice Chairman. Election and duties of two Permanent
Secretaries. Taxonomic subcommittees and their functions. 124

5. Authorization and organization of Judicial Commission. Duties of
Chairman of Judicial Commission. Functions of Judicial Commission. 130

Appendices

A. Transliteration of Greek words into Latin form for use in forma-
tion of scientific names. 137

B. Recommendations relative to choice of spelling between ortho-
graphic variants, or alternative spellings of names and epithets. 143

C. Opinions relating to nomenclature of the bacteria and viruses. 147

D. Lists of conserved and rejected names. 165

Index 173

CHAPTER 1

General Considerations

CHAPTER 1

General ConsideratLons

GENERAL CONSIDERATION 1. The progress of bacteriology can be furthered
by a precise system of nomenclature which is properly integrated with the
systems used by botanists and zoologists and accepted by the majority of
bacteriologists in all countries. The Bacteriological Code applies to bacteria,
related organisms and the viruses. Botanical and Zoological Codes provide
for the nomenclature of certain other microbial groups such as the yeasts
and fungi, algae and protozoa. The special nomenclatural problems of these
groups require cooperation with zoologists and botanists.

ANNOTATIONS*

General Consideration 1. The Botanical Code states (Preamble) :

Botany requires a precise and simple system of nomenclature used

by botanists in all countries, dealing on the one hand with the

terms which denote the ranks of taxonomic groups or units, and on

the other hand with the names which are applied to the individual

taxonomic groups.

No court is competent to enforce the provisions of any nomenclatural

code. Adherence to the provisions of the Bacteriological Code is left

to the individual judgment of the microbiologist. Nonconformity

seems to be due more to lack of knowledge of the Code or to lack of

understanding of its intent than to fundamental disagreement with

the provisions themselves.

There are three official codes of nomenclature, one for each of the
three segments of biology: botany, zoology and bacteriolog\ (includ-
ing virology) . The existence of the three codes is troublesome, for
the microbiologist works with microorganisms whose nomenclature
is determined by the Bacteriological Code (the bacteria and viruses) ,

* These annotations are quite unorticial. They are intended to be helpful in
the interpretation of the Rules and Recommendations, by giving examples and by
noting corresponding Rules and Recommendations to be found in the Botanical
and Zoological Codes.

[3]

4 International Code of Nomenclature

General Consideration 1.

by the Botanical Code (the fungi, algae and sHmc molds) and by the
Zoological Rides (the protozoa) . Fortunately, the more important
rules of the three codes are similar; differences are reasonably clear
but must be understood. The rules and recommendations of this
Bacteriological Code for the most part agree with those of the Botani-
cal Code; wherever there are noteworthy differences between the pro-
visions of the Bacteriological and the Botanical or Zoological Codes,
they are noted in the appended Annotations. Should the rules and
recommendations of the several codes prove to be in conflict and lead
to doubt as to the correctness or legitimacy of a name. General Con-
sideration 1 suggests jjrovisions for coordination through consulta-
tion with the appropriate representative committees of botanists or
zoologists. Botanists, through action of International Botanical Con-
gresses and the International Bureau for Plant Taxonomy and
Nomenclature have organized special committees and subcommittees
to work on problems relating to particular groups, for example the
algae, the fungi, the lichens and the bacteria. Certain microbiological
problems can be referred to such committees for advice or for action.
The Zoological Rules also provide for committees of specialists to
consider and make recommendations relative to problems of the
several subdivisions of the animal kingdom. The Judicial Commis-
sion of the International Committee on Bacteriological Nomenclature
is directed to work actively with the corresponding committees in
zoology and botany on matters of common interest.

A related statement in the International Rvdes of Zoological
Nomenclature is:

The object of the Rules is to provide a system under which the
name of each taxon is unique and distinctive. A primary purpose is
to insure the stability and universal acceptance of names. The
Rules do not trespass on freedom of taxonomic practice.

Bacteria and Viruses 5

General Consideration 2.

GENERAL CONSIDERATION 2. The precepts on which this system of bac-
teriological and viral nomenclature is based are divided into Principles,
Rules and Recommendations.

Principles (Chapter 2) form the basis of the rules and recommendations.
Rules (Chapter 3) are designed

(a) to make effective the principles given in Chapter 2,

(b) to put the nomenclature of the past in order, and

(c) to provide for the nomenclature of the future.

The rules in general are retroactive; names or forms of nomenclature con-
trary tc a rule (illegitimate names or forms) cannot be maintained. Certain
rules authorize appendices and define their scope and authority.
Recommendations deal with subsidiary points, their object being to bring
about greater uniformity and clearness, especially in future nomenclature;
names or forms contrary to a recommendation cannot on that account be
rejected, but they are meant to prevent a repetition of undesirable pro-
cedures of the past. Recommendations are appended to the rules which they
supplement. Notes attached to rules or recommendations are intended to be
explanatory or illustrative and have the same force as the rules or recom-
mendations to which they are appended.

ANNOTATIONS

General Consideration 2. The Principles may be regarded in a
sense as the constitutional background of the rules, recoynynendations,
notes, appendices, lists, etc. They are not rules, but may be useful
in interpretation, helpful in promoting a better understanding
of the rules, and in the solution of problems not covered specifically
by them. In theory at least there should be no conflict between the
rules and the principles.

The Rules are the laws which determine the correctness of names
which have been given in the past and constitute a guide to the
giving of new names. The corresponding rule of the Botanical Code
reads in part:

They are always retroactive except when expressly limited.

The Appendices supplement the rules. In them are listed
Opinions and various groups of conserved and rejected names; they
also include discussions and material too lengthy for convenient in-
corporation directly into the rules or the recommendations.

The Recot7i7ne77dations include the directives of good usage.
Authors show good taste and judgment when they propose names
that conform to recommendations. In some cases the Bacteriological
Code authorizes the correction of names which do not conform to a
particular recommendation. However, the provision that non-con-
formity to a recommendation is not sufficient cause for rejection of
a name should be emphasized.

The Botanical and Bacteriological Codes are in essential agree-
ment in their definitions of Principles, Rules and Recommendations.

6 International Code of Nomenclature

General Consideration 3.

GENERAL CONSIDERATION 3. Provisions (Chapter 4) for emendations of
rules, for special exceptions to rules, and for their interpretation in doubtful
cases have been made through the establishment of an International Com-
mittee on Bacteriological Nomenclature for the International Association of
Microbiological Societies and of its Judicial Commission.

ANNOTATIONS

General Consider atioyi 3. Chapter 4 (p. 123) includes Provisions
made for selection of a large committee with representation from all
countries having microbiological societies which desire representa-
tion. This International Conmiittee on Bacteriological Nomenclature
(or more briefly, the Nomenclature Committee) elects a Judicial
Commission of twelve, to which the two permanent secretaries of the
Committee are added, making a total membership of fourteen. Chap-
ter 4 assigns to the Nomenclature Committee and the Judicial Com-
mission many legislative, judicial and editorial functions. The
Nomenclature Committee was authorized and organized in 1930, the
Judicial Commission in 1939.

The Nomenclature Committee has appointed several subcommit-
tees to consider the problems of certain taxonomic groups.

Botanists, by action of International Botanical Congresses, have
developed a Nomenclature Section, several Nomenclature Committees
and an affiliated International Association for Plant Taxonomy and
Nomenclature.

The zoologists have a corresponding but somewhat different series
of legislative and judicial agencies authorized by the several Inter-
national Zoological Congresses.

See Chapter 4 and its Annotations for details.

CHAPTER 2

Principles

CHAPTER 2

Principles

PRINCIPLE 1. The essential points in nomenclature are

(1) to aim at fixity of names;

(2) to avoid or to reject the use of names which may cause error or
ambiguity or throw science into confusion.

Next in importance is the avoidance of all useless creation of names. Other
considerations, such as absolute grammatical correctness, regularity, or
euphony of names, more or less prevailing custom, regard for persons, etc.,
notwithstanding their undeniable importance, are relatively accessory.

PRINCIPLE 2. In the absence of a relevant rule, or where the consequences of
rules are doubtful, established custom must be followed. In doubtful cases a
resume in which all pertinent facts are outlined should be submitted to the
Judicial Commission for an Opinion.

PRINCIPLE 3. Bacteriological nomenclature and botanical nomenclature are in-
terdependent in the sensa that the name of a bacteiial taxon is to be rejected
if it is a later homonym of the name of any plant taxon. Likewise, nomen-
clature of bacteria and protozoa are interdependent: the name of a bacterial
taxon is to be rejected if it is a later homonym of the name of a protozoan
faxon. Bacteriological nomenclature is independent of zoological nomen-
clature (protozoology excepted): the name of a bacterial taxon is not to be
rejected simply because it is identical with the name of a taxon in the
animal kingdom.

ANNOTATIONS

Principle 1. This principle is similar in phraseology to the state-
ment in the Preamble of the Botanical Code. It is basic to the in-
terpretation of the rules of nomenclature.

The corresponding statement of the Zoological Rules is:

A primary purpose is to insure the stability and universal acceptance

of names.

[9]

10 Infernat'lonal Code of Nomenclature

Principle 2. Principle 3. Annot. cont.

Principle 2. For comment on the format of a request for an Opinion,
see Provision 5 (p. 131) .

Principle 3. The Botanical Code (Principle I) states:

Botanical nomenclature is independent of zoological nomenclature
in the sense that the name of a plant must not be rejected merely be-
cause it is identical with the name of an animal.
Article 45 of the Botanical Code reads in part as follows:

If a taxon is transferred from the animal to the plant kingdom,
its name or names valid* under the International Rules of Zoolo-
gical Nomenclature and validly published in the form provided in
the Botanical Code (except that for algae, validity under the zoologi-
cal rules only is required) shall be automatically accepted as having
been validly published under this Code at the time of its valid
publication as the name of an animal.
The corresponding Article 1 of the Zoological Rules reads:

Zoological nomenclature is independent of botanical nomenclature
in the sense that the name of an animal is not to be rejected simply
because it is identical with the name of a plant. If, however, an
organism is transferred from the vegetable to the animal kingdom,
its botanical names are to be accepted in zoological nomenclature
with their original botanical status; and if an organism is trans-
ferred from the animal to the vegetable kingdom, its names retain
their zoological status for purposes of homonymy.
The Zoological Rules also include the following recommendation:

Avoid introducing into zoology a generic name used in botany.
It would be well to avoid proposal of new names of taxa in bac-
teriology that have been used in zoology.

* The word "valid" in the International Rules of Zoological Nomenclalure is
equivalent to "legitimate" in the International Code of Botanical Nomenclature.

Bacteria and Viruses 7 7

Principle 4.

PRINCIPLE 4. Scientific names of all taxonomic groups (taxa) are usually
taken from Latin or Greek. When taken from any language other than Latin,
or formed in an arbitrary manner, they are treated as if they were Latin.
Latin terminations should be used so far as possible for new names.

ANNOTATIONS

Principle 4. The meaning of the phrase "usually taken from
Latin or Greek" is made clear by prevailing custom. The phrase does
not mean that only those Latin and Greek words as found in the
dictionaries and lexicons may be used, but that new words proposed
as names or epithets may also be coined from the stems of these
words singly or as compounds. It is assumed that the classic tradi-
tion for the forming of new names will be followed.

Appendix A may be used as a guide for the transcription of
Greek into Latin form and for the use of the appropriate Latin
gender endings, as prescribed in the last sentence of this Principle.

This statement is a Principle and not a Rule. The Botanical
Code, from which this Principle was taken, has altered the last sen-
tence from the form of a Recommendation to a definite requirement.
It reads:

Latin terminations are used so tar as possible for ne^v names.
The Zoological Rules state:

The scientific name of an animal must be a word that is either Latin
or Latinised or considered and treated as such in case it is not of
classic origin.

The standard of classical Latin is not to be applied in such a man-
ner as to ignore later developments of the language or to override
considerations of scientific accuracy, uniformity, intelligibility or
practical usefulness.
This Principle applies to names of genera and subgenera and to
specific and subspecific epithets. In general, it is clearly intended that
a name taken from a language such as Greek shotild be transliterated
in accordance with classic usage, and that the word be placed in the
appropriate Latin declension with Latin endings. Each of the bio-
logical codes of nomenclature has added some rules and recommenda-
tions to supplement those of classic usage for situations not en-
visaged in classic Latin.

Section 1 (Chapter 3) , Rules 1-9 on "Naming of Taxa of Various
Ranks" and Section 7 (Chapter 3) , Rules 27-28 on "Orthography
and Gender of Names" with their Recommendations, together with
.Appendices A and B, spell out in some detail the interpretation of
this Principle 4. The types of problems (with examples) encountered
are discussed in the several annotations.

72 International Code of Nomenclature

Principle 5.

PRINCIPLE 5. Nomenclature deals with:

(1) The terms which denote the categories of taxa (taxonomic groups or
units, such as species, genus, family) and the relative ranks of these
categories.

(2) The names which are applied to the individual taxa (taxonomic
groups), such as Bacillus subtilis. Streptococcus, Spirillaceae, Spiro-
chaetales.

NOTE 1. The use of the word taxon (plural taxa) is appropriate for the term

taxonomic group or the word group wherever used in the sense of taxonomic

group in this Code. The word group has as its preferred use in bacteriology

that indicated in Recommendation 8a.

NOTE 2. The word name, unless otherwise indicated, means a name which

has been validly published, whether legitimate or illegitimate.

A legitimate name or epithet is one that is in accordance with the rules.

An illegitimate name or epithet is one that is contrary to the rules.

The correct name of a taxon with a particular circumscription, position, and

rank is the name which must be adopted for it under the rules.

A name is effectively published when its publication is in accordance with

Rule 11.

A name is validly published when its publication is in accordance with Rules

12, 13 and 14.

ANNOTATIONS

Principle 5. The Botanical Code has practically identical word-
ing. The Zoological Code neither defines nomenclature nor notes the
differentiation of terms denoting categories and names of taxa,
though there is in this Code implicit recognition of the existence of
distinctions.

The names and relative ranks of the several categories of taxa
are given in Principle 7 (p. 13) .

A^ote 1. The statement in the Botanical Code reads:
Taxonomic groups of any category will, in this Code, be referred
to as taxa (singular, taxon) .
The fourteenth International Congress of Zoology at Copenhagen,
1953, approved the introduction of the expression "taxon" to repre-
sent the concept "taxonomic unit." "The expression is to be used
wherever appropriate, throughout the Zoological Code."

Note 2. The definitions of the several phrases as given in the
Bacteriological Code correspond to those of the Botanical Code.

The Zoological Code does not recognize certain of the phrases
of Note 2, and uses a different terminology for others, particularly
the following:

An available name is one which is "sufficient nomenclaturally" (as
contrasted with "taxonomically") . Apparently this corresponds in
part to the phrase "legitimate name" of Note 2.

A valid name is one which is "'sufficient taxonomically" (as con-
trasted with "nomenclaturally") .

The phrase "duly published" is apparently used in much the same
sense as "validly published" in the Bacteriological Code.

Bacteria and Viruses 73

Principles 6 and 7.

PRINCIPLE 6. The rules and recommendations of bacteriological nomenclature
apply to all bacteria, recent and fossil, with certain distinctly specified ex-
ceptions.

ANNOTATIONS

Principle 6. The corresponding statement of the Botanical Code
reads:

The rules and recommendations of botanical nomenclature apply
throughout the plant kingdom, recent and fossil, with certain dis-
tinctly specified exceptions.

Note. In general the International Code of Botanical Nomenclature
applies also to Bacteria. However, many special provisions are
needed for this group and for this reason a special International
Code of Bacteriological Nomenclature has been provided by the
International Microbiological Congress. (See Jour. Gen. Micro-
biology 3 (3) :444. 1949) .

PRINCIPLE 7. The terms which denote the rank of taxonomic groups (taxa)
are defined as follows:

(a) Every individual is treated as belonging to a number of categories of con-
secutive rank and consecutively subordinate; of these the species is the basic
one. The principal categories in ascending sequence are species (species), genus
(genus), family (familia), order (ordo), class (classis), divisi:»n (divisio). In some
genera the rank subgenus may be distinguished, and in some families the
rank tribe.

(b) In many species, subspecies or varieties are distinguished; in some cases
subdivisions of a species such as strains, groups, serotypes, variants, phases
and others may be recognized.

(c) If a greater number of intermediate categories (ranks) are required, the
terms for these subdivisions are made by adding the prefix "sub-" to the terms
denoting the ranks. Thus, subfamily denotes a rank between a family and a
tribe, subtribe a rank between a tribe and a genus, etc.

(d) The definition of each of these categories (ranks) varies, up to a certain
point, according to individual opinion and the state of the science; but their
relative order, sanctioned by custom, must not be altered. No classification is
admissible which contains such alterations.

ANNOTATIONS

Principle 7. The categories of taxa that are recognized in the
Bacteriological Code may be summarized as follows:

1. (Division) (Divisio) ' 9. (Tribe) (Tribus)

2. (Subdivision) (Subdivisio) 10. (Subtribe) (Subtribns)

3. Class (Classis) 11. Genus (Genns)

4. (Subclass) (Subclassis) 12. (Subgenus) (Subgenus)

5. Order (Ordo) 13. Species (Species) '

b. (Suborder) (Subordo) (14a. (Subspecies) (Subspecies)

7. Family (Familia) jl4b. (Variety) (Varietas)

8. (Subfamily) (Subfaniilia) 15. Individual (Individuurn)

14 International Code of Nomenclature

Principle 7. Annot. cont.

The above categories of taxa are recognized also in the Botanical
Code. Certain others arc recognized as admissible. One may insert
between Subgenus and Species the categories Section (Sectio) , Sub-
section (Subsectio) , Series (Series) and Subseries {Siibseries) .

The Bacteriological Code also recognizes certain subdivisions of
species (Recommendation 8a 1-6) such as Strain, Serotype, Group,
Phase, Forma Specialis, Variant, Stage and State. These are termed in-
iVasubspecific forms and are to be regarded as categories of taxa.

The Zoological Code has definitely recognized relatively few cate-
gories. A series of groups of categories or ranks is in process of de-
velopment at the present time (1957) . The group of categories from
Subphylum upwards is to be called the "Phylum Group," that from
Suborder to Superclass inclusive is the "Order/Class Grotip," be-
tween the Generic Group and the Order/Class Group is the "Family
Group," followed by the "Generic/Subgeneric Group," and the
"Species/Subspecies Group." The category variety is not recognized.

The complexity of the problem of subdividing species has led the
zoologists to recognize "infrasubspecific" names for any "form of a
species other than a subspecies." Many of the problems of nomen-
clature in this group of infrasubspecific forms resemble those met
in bacteriology.

The hierarchy of categories of taxa has not been as definitely
fixed in zoology as in bacteriology and in botany. This situation
has come about in part by the development of special precedents and
even separate nomenclatural codes in the pa^t by zoological workers
(as the entomologists) in large and special groups.

Bacteria and Viruses 15

Principles 8 and 9.

PRINCIPLE 8. The primary purpose of giving a name to a taxonomic group
is not to indicate the characters or the history of the group, but to supply
a means of referring to it.

ANNOTATIONS

Principle 8. The Botanical Code adds to this statement: "and to
indicate its general taxonomic position."

In some cases names are long and cumbersome because an author
conceived it his duty to include as full a description of the taxon as
practicable. It is desirable, but not necessary, that names be ap-
propriate and descriptive. A generic name such as Deazotonitranitria-
zobacterium Ambroz 1913 was an attempt to indicate the fact that
bacteria of this genus are small rods capable of producing free nitro-
gen from nitrates and nitrites. The tendency to form descriptive
epithets, if not too long, is laudable and is definitely advised
(see Recommendation 6b) . But if the advice is not taken, and
a name (particularly a specific epithet) is given which is regarded
by a subsequent author as nondescriptive or even unsuitable, such
name is not to be changed for this reason; it still serves the useful
purpose of supplying "a means of referring to it."

PRINCIPLE 9. Each order or taxon of lower rank with a given circumscription,
position and rank can bear only one correct name, the eailiest that is in ac-
cordance with this Code of Nomenclature. Provision is made for exceptions
that have been approved for inclusion in the list of nomine conservonda.
In subgenera, genera, and groups of higher rank to order inclusive, the correct
name is the earliest name published, provided that this is in conformity with
the rules of this Code. In species the correct name is the binary and in sub-
species the ternary combination of the generic name with the earliest available
legitimate epithet validly published with the same rank.

ANNOTATIONS

Principle 9. This principle is the basis in the Rules for Sections 1
(Naming of Groups of Various Ranks) , 3 (Publication of Names) , 5
(Changes in Names as a Result of Segregation, or Union of Taxa or

Changes in Rank of Taxa) and 6 (Rejection and Replacement of

Names) .

16 International Code of Nomenclature

Principle 10.

PRINCIPLE 10. Bacteriologists are urged not to change the name of a taxon
without serious motives, based either on more profound knowledge of facts
or on the necessity of giving up a nomenclature that is contrary to the Rules
of this Code.

ANNOTATIONS

Principle 10. The corresponding Article 17 of the Botanical Code
(Seventh Botanical Congress, Stockholm, 1952) was included as a
Rule with a Recommendation which read:

Art. 17. No one may change a name (or combination of names)
without serious motives, based either on more profound knowledge
of facts or on the necessity of giving up a nomenclature that is con-
trary to the rules of this Code.

Recommendation 17A. Changes in nomenclature should be made
only after adequate taxonomic study.

The Nomenclature Section of the eighth International Botanical
Congress (Paris, 1954) agreed that the above Article 17 should not
be included in the rules but transferred to Chapter I, Art. 2, under
"General Considerations, Guiding Principles and Definitions", and
that consideration should be given by the Editorial Committee to the
following substitute wording:

No one can change a name (or a combination of names) without
serious motives based either on some adequate taxonomic study or
on the necessity of giving up a nomenclature that is contrary to the
rules of this Code.

One may conclude that the phrase "based on more profound
knowledge of facts" may well be interpreted to mean "based upon
some adequate taxonomic study."

The Zoological Code does not directly rule on the above problem.

Bacteria and Viruses 17

Principle 11.

PRINCIPLE 11. The application of the names of taxonomic groups (taxa) is
determined by means of nomenclatural types. A nomenclatural type is that
constituent element of a taxon to which the name of the taxon is perma-
nently attached.

NOTE. The nomenclatural type is not necessarily the most typical or repre-
sentative element of a taxon. It is merely that element with which the name
of a taxon is permanently associated.

ANNOTATIONS

Principle 11. The meaning and significance of this principle defining
the phrase nomenclatural type are developed by Rule 9 (j). 53) .
For example, the species Bacillus subtilis has been designated as the
nomenclatural type of the genus Bacillus. Whenever a genus Bacillus
is recognized by an author, he must include within it the species
Bacillus subtilis. One might even define the genus Bacillus as con-
sisting of the type species, Bacillus subtilis, together with such other
species as are placed with it because presumably sufficiently closely
related (congeneric) .

The corresponding statement in the Botanical Code of 1952 (Article
18) is essentially the same, with the exception of the second sentence,
which reads:

A nomenclatural type (typus) is the constituent element of a taxon

to which the name of the taxon is permanently attached, whether

as an accepted name or as a synonym.
The type concept is outlined in Articles 18, 19, 21 and 22.

The Zoological Code outlines the type concept in much greater
detail than does either the Bacteriological or Botanical Code.

18 Infernational Code of Nomenclature

Principle 12.

PRINCIPLE 12. A name of a taxonomic group has no status under the Rules
and no claim to recognition by bacteriologists unless it is validly published.

ANNOTATIONS

Principle 12. Section 3 ot the Bacteriological Code, Rules 10-14
(pp. 59-71) outline the requirements for \alid publication.

The Botanical Code gives essentially the same Rules and Recom-
mendations in its Articles .S9-54, but in greater detail. The differences
between the codes will be noted in the Annotations on pp. 59-71.

The Zoological Code likewise has a long list of Rules and Recom-
mendations relative to publication which are noted also in the An-
notations.

CHAPTER 3

Rules of Nomenclature

With
Recommendations

CHAPTER 3

Rules of Nomenclature
With. Recommendations

Section 1

Naming of Taxonomic Groups (Taxa) of Various Ranks

RULE 1. The names of all taxonomic groups (taxa) above the rank of genus
are substantives or adjectives used as substantives, of Greek or Latin origin,
or Latinized words, in the plural number.

ANNOTATIONS

Rule 1. Basically the names of all taxa from genus to order inclusive
are plural adjectives modifying the word plantae. They are now
recognized and treated as nouns. Rule 4 designates special endings
that are to be used as suffixes in the formation of names of these
higher taxa. The plural adjectival endings in general have the mean-
ing of "like" or "resembling."

RULE 2. Names of all taxonomic groups (taxa) above the rank of family are
taken preferably from a combination of characters covering the nature of
the taxa as closely as possible, or from a single character of outstanding
importance.

RECOMMENDATION 2a. Names of new orders and suborders should prefer-
ably be based upon the name of the type genus of a contained family.

ANNOTATIONS

Rule 2. The corresponding Article (16) in the Botanical Code
reads:

The principles of priority and typification do not apply to names
of taxa above the rank of order.

Recommendation 16a of the Botanical Code suggests that certain
endings for taxa above the rank of order be used. It states:

(a) The name of a division is preferably taken from characters in-
dicating the nature of the division as closely as possible; it should
end in -phyta, except when it is a division of Fungi, in which case
it should end in -mycota. Words of Greek origin are generally pre-
ferable.

The name of a subdivision is formed in a similar manner; it is dis-
tinguished from a divisional name by an approprate prefix or suf-
fix or by the ending -phytina, except when it is a subdivision of
Fungi, in which case it should end in -mycotina.

(b) The name of a class or of a subclass is formed in a siniiliar
manner, and should end as follows:

1. In the Algae: -phyceae (classes) and -phycidae (subclasses);

2. In the Fungi: -mycetes (classes) and -mycetidae (subclasses) ;

3. In the Cormophyta: -opsida (class) and -idae (subclasses).

In bacteriology little or no use is made of names of taxa above that
of class.

[21]

22 International Code of Nomenclature

Rule 3. Chapter 3. Sect. 1. Naming of Taxa

RULE 3. Names of taxonomic groups (taxa) between suborder and genus are
formed by the addition of the appropriate suffix to the stem of the name of
the type genus.

ANNOTATIONS

Rule 3. The Bacteriological and Zoological Codes have equivalent
statements. The Botanical Code has no over-all formulation but
states the case for families as follows:

The name of a family is a plural adjective used as a substantive
formed by adding the suffix -aceae to the stem of the name of its
type genus or of a synonym of this name, even if illegitimate.
The Zoological Rules state:

When the name of the type genus (or its terminal portion) is a
word of Greek or Latin origin, its genitive termination is to be re-
placed by the appropriate Family-Group temiination (e.g. "-idae"
for family, "-inae" for subfamily) . If incorrectly formed when first
published, such a group name is to be automatically corrected. \
Schedule to the Rules is to illustrate the correct method of forming
such names.

Where the name of the type genus is not of Greek or Latin origin,
or is an arbitrary combination of letters, the name first published
for that Family-Group unit with the appropriate termination is to
be accepted.

An author, when forming a Family-Group name based on a word
not of Greek or Latin origin, should select such portion of the name
of the type genus as, with the appropriate tenuination, will reveal
the relation between the generic name and that of the Family-
Group unit, and will give the name so formed the simplest and
most euphonious form compatible with that relationship.
The Bacteriological and Botanical Codes agree that the appropriate
suffix of the taxonomic rank is to be added to the stem of the name
of the type genus (or, in botany, alternatively to the name of a syno-
nym of the type genus) .

The Zoological Code directs that the suffix is to replace the geni-
tive ending of the name of the type genus. Statements of the Zoologi-
cal and Bacteriological Codes are intended to be equivalent.

It will be noted that the Zoological Code is to provide a Schedule
to be appended to the Rules to illustrate what changes are to be
made in the name of the type genus before adding the suffix indi-
cating the rank of the new taxon.

Such a Schedule was proposed for the Botanical Code by Ponce
de Leon and Alvarez (1 axon 5:45. 1954) but was rejected.

Bacteria and Viruses 23

Rule 3. Chapter 3. Sect. 1. Naming of Taxa

Determination of the spelling ol stems or combining forms to be
used with appropriate suffixes is primarily a problem of orthography,
and the discussion is delayed to the annotations of Section 7 on
Orthograph\ :iud Gender of Names (Rule 27, p. 99ff.) , q.v.

24 International Code of Nomenclature

Rule 4. Chapter 3. Sect. 1. Naming of Taxa

RULE 4. Names of taxonomic groups between subclass and genus have suf-
fixes to fix the taxonomic rank. The suffix for orders is -ales, for suborders
-ineoe, for families -oceoe, for subfamilies -oideoe, for tribes -eae and for
subtribes -inoe.

ANNOTATIONS

Rule 4. The suffixes specified in the Bacteriological Code for forma-
tion of names of higher taxa between subclass and genus and those
authorized by the Botanical Code are identical. The Zoological Code
specifies two suffixes only: all family names must end in -idae and
subfamily names in -inae. No endings to be used for other higher
taxa have been officially recognized.

The suffixes used in both Bacteriological and Botanical Codes
are all plural feminine adjective endings. The singular forms of
these suffixes are -alls, -inea, -oidea, -ea and -ina. The suffix oidea is de-
rived from the Greek, -ina may be either a Greek or a Latin suffix,
the others are Latin adjectival suffixes. All as here used have the
connotation of "like" or "resembling" or "having the characteristics
of." These endings are in the feminine plural as they in theory
modify the plural Latin noun Plant ae. The family name Spirillaceae
literally has the meaning of "spirillaceous plants," that is, plants
(organisms) resembling those of the genus SpiriUit7n. However, in
modern nomenclature the names of taxa above genus are treated as
plural substantives (nouns) and not as adjectives. Further discus-
sion of the formation of names of higher taxa will be found in the
annotations in Section 7 under "Orthography and Gender of Names"
(p. 99ff .) .

Occasionally (rarely) one finds the name of one of the higher
taxa used in the singular. One may, for example, designate an or-
ganism as a Spirillacea meaning that it is one of the species belong-
ing to the family Spirillaceae. This is done not infrequently in
Botany; the phrases "this is a Rosacea" or "a new Rosacea" are briefer
than "this belongs to the Rosaceae" or "a new species of Rosaceae."
Apparently this usage has been confined almost entirely to the singu-
lar of family names; however, such use of a family name in the
singular is rare in microbiology.

Authors sometimes change the spelling of names of higher taxa
to conform to the characteristic of the vernacular in which they write.
In French the family name Spirillaceae may change from its Latin
form to Spirillacees and in German to Spirillaceen (or more fre-
quently to Spirillazeen) . Trevisan used as a title for an important
taxonomic brochure "I generi e le specie delle Batteriacee." Billet
used the phrase "d'une nouvelle bacteriacee marine." This is not

Bacteria and Viruses 25

Rule 4. Sect. 1. Naming of Taxa

customary in English; one docs not often find a usage such as spiril-
laceas. Such words with the modified spellings are no longer Latin,
they are vernacidar, and are not the names authorized by the inter-
national agreements of the several codes of nomenclature. They may
be justified in non-scientific writing but cause difficulty when, as
has happened, new names of higher taxa such as families are pro-
posed in the vernacular spelling. Does such a word have standing
in nomenclature when not spelled as a Latin word? Should such
words be discarded as not validly published and replaced, or should
the incorrect spellings be regarded as unintentional error and cor-
rected? No formal Opinion has been rendered. It may be that the
legitimacy of such vernacular names of higher taxa should be con-
sidered individually.

26 International Code of Nomenclature

Rule 5. Chapter 3. Sect. 1. Naming of Taxa

RULE 5a. Names of genera and subgenera are substantives (or adjectives
used as substantives) in the singular number and written with an initial
capital. The names may be taken from any source whatever and may even
be composed in an arbitrary manner. They are treated as Latin substantives.
RULE 5b. Generic names and subgeneric names are subject to the same rules
and recommendations, and from a nomenclatural standpoint they are co-
ordinate.

RULE 5c. If a genus is divided into subgenera, one of the subgenera (that which
includes the type species of the genus) must bear the same name as the genus.
NOTE. Citations of authors are governed by Rule 16.

ANNOTATIONS

Rule 5. Genera and subgenera. Rule 5a is much like that of the
Botanical Code.

Ride 5 is similar in intent to several articles in the Zoological
Rules, which provide that:

1. A generic name must consist of a single word, simple or com-
pound, written with a capital initial letter, and employed as a
substantive in the nominative singular.

2. Generic and subgeneric names are subject to the same rules and
recommendations, and from a nomenclatural standpoint, they
are co-ordinate, that is, they are of the same value.

3. A generic name becomes a subgeneric name when the genus so
named becomes a subgenus, and vice versa.

4. If a genus is divided into subgenera, the name of the typical
subgenus must be the same as the name of the genus.

Rule 5 makes it clear that if a genus is found to contain two or
more readily differentiable groups of species, each group may be
recognized as a subgenus. One of the subgenera, that which includes
the type species, must have the same name as that of the genus. If
it is desired to recognize each of the groups of species as a distinct
genus, the subgeneric names then become generic names.

Conversely, when two or more genera are luiited, the names of
the genera become the names of subgenera if subgenera are recog-
nized.

The Paris (1954) revision of the Botanical Code has modified the
treatment of taxa between genus and species. The pertinent Article
(21) reads in part:

The name of a subdivision of a genus is a combination of a generic
name and a subdivisional epithet connected by a term (subgenus,
section, series, etc.) denoting its rank.

The epithet of a subgenus or section must not be fonned from the
name of the genus to which it belongs by adding the ending -aides
or -opsis.
1 he Botanical Recommendation 21 A reads:

For a subgenus and a section the epithet is usually a substantive
resembling the name of a genus or repeating the name of the genus
itself.

Bacteria and Viruses 27

Rule 5. Sect. 1. Naming of Taxa

For a subsection and a lower subdivision of a genus the epithet is
preferably a plural adjective agreeing in gender with the generic
name and written with a capital initial letter.

The treatments of subdivisions of a genus are distinctly different
in Botany and in Bacteriology. These differences may be summarized:

1. Bacteriology recognizes the subgenus as the only subdivision
between genus and species. Botany recognizes several, in-
cluding subgenus, section, subsection, series and subseries.

2. Bacteriology and Zoology regard a subgeneric name as subject
to the same rules and recommendations as a generic name.
Botany has a series of special rvdes: the subgenus and genus
are 7iot co-ordinate nomenclaturally.

3. Bacteriology agrees with Zoology in regarding names of all
subgenera, genera and higher groups as uninomial. Botany
regards the name of a subgenus as an epithet related to or
modifying the generic name.

4. Bacteriology and Zoology definitely regard the subgenus as a
taxon. Whether the subgenus in Botany is a taxon is problem-
atical. It is difficiUt to equate epithet and taxon.

5. Bacteriological and zoological rules forbid the use of the same
subgeneric name in different genera. The name of a subgenus
may be the same as that of the genus in which it is included,
but may not be a homonym of the name of another genus. In
Botany the subgeneric epithet in a genus may duplicate the
name of another genus or of an epithet of a subgenus in
another genus.

Names of Genera. Although Rule 5 states that almost any combi-
nation of letters may be proposed and used as a generic name; authors
should hold in mind the directive of Principle 4 which states that
names of taxa are usually taken from Latin or Greek, but if taken
from any other language or formed in an arbitrary manner they are
to be treated as Latin, and Latin terminations should be used so far
as possible. In other words, while any combination of letters may
be used, certain methods of coining generic names are much to be
preferred. One may well observe the Linnaean injunction that the
most appropriate and satisfactory generic name is one derived from
the Greek, usually the result of joining appropriate Greek stems to
form a new word.
Examples of generic names,

With a single Greek stem: Bacterium, Clostridium, Proteus,
Plectridiiim.

Compounds with two Greek stems: Actinomyces, Aerobacter,
Chondromyces, Corynebacterium, Streptococcus, Haemophilus, Tre-
ponema, Thiothrix, Pseudomonas.

With a single Latin stem: Vibrio, Spirillum, Sarcina, Bacillus,
Cloaca.

28 International Code of Nomenclature

Rule 5. Sect. 1 . Naming of Taxa

Compounds with two Latin stems: Putribacilius, Lactobndllus,
Fu.slformis.

Latinized names of persons: Borrelia, Kurthia, Gaffkya, Esche-
richia, Erwinia, Gallionella, Pasteurella, Klebsiella, Eherthella.

Latin-Greek hybrid names: Brevibacterium, Flavobacterium,
Actinobacilhis, Acetobacter, Acetomonas.

Names of Subgejiera. Names of subgenera, as noted, are formed
exactly as are the names of genera. When included in the name of a
species, the subgeneric name is placed in parentheses between the name
of the genus and the specific epithet. If one recognizes two subgenera
in the genus Bacillus, one (that including the type species of the
genus) must bear the same name as the genus. One would then
write Bacillus (Bacillus) siibtilis and Bacillus (Aerobacilhis) poly-
niyxa to designate species of two subgenera differentiated on the
basis of gas production. The name of a subgenus is not combined
with a specific epithet independently of the generic name; the use
of Aerobacillus poly my xa indicates recognition of Aerobacillus as a
generic name. The requirement that one subgenus in a genus must
bear the same name as the genus (Bacteriological Code 1947) would
make necessary the revision of Pederson's (1945) recognition in the
genus Lactobacillus of three subgenera: Thennobacterium, Strepto-
bacterium and Betabacterium; the one containing the type species
should be called Lactobacillus.

Use of the generic name. The commonest use of a generic name
is in combination with a specific epithet to form a binary combination
(the scientific name of a species) . The generic name Bacillus with
the specific epithet subtilis gives Bacillus subtilis. But the generic
name is also commonly used without designation of species. There
is some confusion in the literature of bacteriology and some evidence
of differences of opinion as to such independent use of a generic
name. The generic name singly is often used in the sense of "this
species of" the genus, or even "this individual" or "this strain" of a
species. Custom throughout biology has apparently approved this
usage. One may, therefore, write appropriately "this Salmonella."
The context determines whether reference is to a particular species
of the genus or to an individual strain of a species. The generic
name is frequently used when it is desired to avoid allocating an
isolate to a particular species; the statement, "This Salmonella was
isolated from an egg," merely places the strain in the appropriate
genus.

There is some difference of opinion as to the use of a generic
name in the plural. The Bacteriological Code, as well as others,
states that there can be only one correct name for a genus, and that
no other genus can legitimately have the same name. It has been
urged that inasmuch as there can be but one genus correctly named
Salmonella, the use of the plural Salmonellae would imply the exist-
ence of more than one genus Salmonella^ which, mider the rules, is

Bacteria and Viruses 29

Rule 5. Sect. 1. Naming of Taxa

not allowable; the use of the plural, it is contended, is therefore
inadmissible. However, the plurals of generic names have been, and
are being, used by nomenclators. Such use has been commonly ac-
cepted. The argument for such use may be phrased as follows. Each
species has a name made vip of two words, the name of the genus
followed by a specific epithet. If it is permissible to write "this
Salmonella" meaning a representative of a species of the genus
Salmonella, similarly it should be permissible to write Salmonellae for
two or more species or strains belonging in the genus Salmonella.
This use is well established with the meaning "species of" or "ex-
ample of" or "members of" the genus Salmonella.

Sometimes a generic name in the singular form is used incorrectly
with a plural meaning; for example, in the statement "Salmonella
are abundant in sewage." The argument for this usage is that there
are many species of Salmonella, hence this generic name may be re-
garded as a collective noun, as the word "committee" in English,
and hence may be used either in a plural or singular sense. The
argument is questionable. There is no warrant for the use of the
singular form of a generic name in the sense of "the several species
of the genus Salmonella" or "strains of the genus Salmonella."

Is it advisable that generic names be accepted in modern lan-
guages and the spelling and usage be made to conform to the ver-
nacular?

Two recommendations of the Botanical Code are pertinent:

When writing in modern languages botanists should use Latin sci-
entific names or those immediately derived from them in preference
to names of another kind or origin (popular names) . They should
avoid the use of the latter unless these are very clear and in com-
mon use.

Every friend of science should oppose the introduction into a
modern language of names of plants which are not already there,
unless they are derived from Latin botanical names by means of
some slight alteration.

When a generic name is used repeatedly in general or nontaxonomic
publications, custom has permitted, in fact, encouraged, the use of the
vernacular version of the name. In English the use of the same
spelling is ustially advisable, but without capitalization and without
italicizing. One has the choice of use of the vernacular or the Latin
plural endings. The latter are often preferable to avoid unsatisfactory
use of plurals ending in -s when the Latin name ends in -s. Certainly
"lactobacilli" is a preferred spelling, but one may use "salmonellas"
or "salmonellae."

Orthography of generic names. Annotations relative to formation
of generic names will be found under Section 7 (Orthography and
Gender of Names, Rule 27) and in the several appendices.

30 International Code of Nomenclature

Recommendation 5a. Sect. 1. Naming of Taxa

RECOMMENDATION 5a. Bacteriologists who are forming new generic or
subgeneric names should attend to the following recommendations:

(1) Not to make names very long or difficult to pronounce.

(2) To make names that have an agreeable form readily adaptable to
the Latin tongue.

(3) Not to dedicate genera to persons quite unconnected with bacteri-
ology or at least with natural science or to persons quite unknown.

(4) To avoid the use of adjectives as nouns.

(5) Not to make names by combining words from different languages
(nomina hybrida).

(6) To give a feminine form to all personal generic names, whether they
commemorate a man or a woman.

ANNOTATIONS

Recommendation 5a (1) . Clements (1902) suggested that generic
names should not be longer than six syllables. There are a few that
are longer; but none of these has proved troublesome. Among the
latter are Corynebacterium and Propionibacteriiim.

Some very long generic names have been proposed in the past, but
such proposals usually have been defective in publication, and it has
not been necessary to consider their legitimacy.

One reason for recommending the choice of relatively short generic
names is that from them may be formed the names of higher taxa
(as families, etc.) by the addition of appropriate suffixes, increasing
their length by one, two or even three syllables. The family name
derived from Actinomyces (5 syllables) is Actinomycetaceae (8 syl-
lables) . A family name based upon Propionibacterium would have
ten syllables (Propionibacteriaceae) .

The Zoological Code recommends that new generic and sub-
generic names should be short and "from the Latin standpoint,
euphonious."

A Zoological Recommendation lists twelve classes of words that
may be used as generic names:

1) Greek substantives transliterated by the Latin rules. 2) Compound
Greek words. 3) Latin substantives. 4) Compound Latin words. 5)
Greek or Latin derivatives expressing comparison, diminution,
possession or resemblance. 6) Mythological or heroic names (with
Latin termination) . 7) Proper names used by the ancients. 8)
Modern surnames with a suffix denoting dedication. 9) Names of
ships treated either like mythological names or as modern sur-
names. 10) Words of non-classic origin. 11) Words formed by an
arbitrary combination of letters. 12) Names formed by anagram.

Recoynmendation 5a (2) . Many names considered difficult because
not readily adapted to the Latin come from words transliterated

Bacteria and Viruses 31

Recommendation 5a. Sect. 1. Naming of Taxo

from languages employing an alphabet other than Latin. Such, for
example, are the Arabic and Slavic languages.

Recommendation 5a (3). In general bacteriologists have adhered to
the recommendation not to make inappropriate dedications. There
are very few generic names of bacteria derived from the names of
persons who were not concerned with microbiology.

Recommeyidation 5a (4) . This recommendation, not to use adjec-
tives as generic names, has been ignored in a few instances. For
example, Castellani and Chalmers placed Bacillus faecalis alcaligenes
Petruschky 1896 in a new genus which they named Alcaligenes
("alkali-producing") . "When used as a generic name it is treated as a
substantive ("that which forms alkali") . Among names proposed
for genera and having an adjectival form are Encapsidntus Castel-
lani and Chalmers, Dysenteroides Castellani and Chalmers, Fusiformis
Hoelling and Albofacieus Castellani and Chalmers.

Recommendation 5a (5) . Generic names formed by combining stems
from different languages are common in microbiology, particularly
names which combine stems from Latin and Greek. This recom-
mendation is one of the oldest in biological nomenclature. Lin-
naeus voiced disapproval of nomiyia hybrida. But violations of the
spirit of the recommendation are numerous and continuing. There
are many reasons for apparent disregard for this recommendation.

For example, an author in search of a name for a new genus may
wish to indicate that his organism has some of the characters of the
genus Bacillus but is not a true Bacillus. He coins the new generic
name Pseudobacillus, a false Bacillus. But the name is a hybrid and
hence undesirable; the first stem pseud- is Greek and bacillus is Latin.
The author could have chosen the Latin word falsus also meaning
false and could have coined Falsibacillus in which both component
stems are Latin. The Latins themselves occasionally used fals- as
the first component of a compound word. Even more frequently a
Latin prefix is the first component of a compound in which the
second is Greek. An author wishes to publish a generic name with
the meaning "short rod." He knows that the Greek word bac-
terium means rodlet. The Latin word for short is brevis, the Greek
is brachys. He chooses Brevibacterium and sponsors a Latin-Greek
compound rather than the etymologically more correct Brachybac-
terium. It has been argued, and perhaps with justification, that a
Greek word such as bacterium, when it becomes a generic name
(Bacterium) , is under the rules of all codes to be treated as Latin
(see Principle 4 of this Code.) If the generic name is to be regarded
and treated as Latin, why should not Latin prefixes be used? With

32 International Code of Nomenclature

Recommendation 5a. Sect. 1. Naming of Taxa

this interpretation, Breinbnctrrium would not mean the "short rod-
let" but the "short Bacterium."

However, hybrid generic names, if validly published, have the
same status as though they were correctly formed.

The Zoological Rules recommend that a Greek word should not
be combined with a Latin word in forming a generic name.

Recoynmenddtion 5a (6) . In Latin the names of most plants (not
all) are feminine. Obviously, if all generic names in botany or bac-
teriology were feminine some problems of nomenclature would be
simplified. Later it will be noted that it is recommended that generic
names formed from the names of persons, whether men or women,
should end in -a or -ia and be treated as feminine nouns.

This recommendation has not always been followed; there have
been proposed as bacterial generic names such derivatives as Wesen-
hergiis Castellani and Chalmers, and MaciutosJiiUus Heller.

The Zoological Code does not recommend that personal names
be Latinized into feminine generic names, but suggests the use of
endings which are masculine, feminine or neuter. In practice, how-
ever, zoologists have usually preferred to put generic names derived
from patronymics in feminine form.

The Botanical Code includes several additional recommendations
of interest, namely:

(a) To use Latin terminations inasfar as possible.

(e) To indicate, it possible, by the formation ot ending ot the name

the affinities or analogies of tfie genus.
(g) Not to use a name similar to or derived from the epithet of one
of the species of the taxon.

Bacteria and Viruses 33

Chapter 3. Sect. 1. Naming of Taxa

RULE 6. The name of a species is a binary combination consisting of the
name of the genus followed by a single specific epithet. If an epithet con-
sists of two or more words, these must either be united or hyphenated.
NOTE. A specific epithet consisting of two or more words not joined by
hyphens when originally published is not to be rejected, but when used the
words are to be hyphenated or joined.
Specific epithets are:

(1) Adjectives, which must agree grammatically with the generic name.

(2) Substantives, in the nominative, in apposition with the generic name.

(3) Substantives in the genitive.

A specific epithet may be taken from any s=u:ce whatever and may even
be composed arbitrarily. Within the same genus, no two species names
may bear one specific epithet.

NOTE. The term "epithet" as used here implies a single descriptive word
or a single descriptive phrase consisting of two or more words. If the author
of the name of a species proposed an epithet consisting of two or more
words, but without hyphenating or joining, the form is to be corrected, but
without prejudice.

ANNOTATIONS

Rule 6. The first two sentences of this rule agree with the statement of
the Botanical Code. The word "combination" as here used is defined in
the Botanical Code: "The name of a taxon below the rank of genus,
consisting of the name of a genus combined with one or more epi-
thets, is termed a combination." The statement relative to the kinds
of specific epithets agrees also with that of the Zoological Code. The
latter, however, includes in the rule directives as to the formation of
specific epithets indicating dedication to one or more persons (see
Rule 27) .

Discussion of formation of specific epithets from names of per-
sons is to be found in Chapter 3, Sec. 7, Rule 27 and its Recom-
mendations.

The nafne of a species. The rule is clear. The name of a species
is a binary combination of the name of the genus followed by a
specific epithet. The specific epithet is not the 7iaj7ie of the species;
it should not be used singly to designate a species.

The Zoological Code emphasizes this point:

The scientific designation of animals is uninominal for subgenera

and all higher groups, binominal for species, and trinominal for

subspecies.

There is, however, confusion in biological literature due to lack
of agreement among those concerned with taxonomy and nomen-

34 International Code of Nomenclature

Rule 6. Chapter 3. Sect. 1. Naming of Taxa

clature as to the appropriate designation ot the second component
of the species name, as well as to the designation of the name of a
species. In bacteriology and botany this second component is termed
the specific epithet. The directive imder the Zoological Rule is quite
different. The Paris Zoological Congress (1948) agreed that the
"term 'specific name' should be reserved to denote the binominal
combination which constitutes the scientific name of a species." This
statement conforms to usage in bacteriology and botany. It was also
agreed at Paris that the

expression "trivial name" (based on early Linnaean usage) should
be adopted to denote the second part of the binominal combi-
nation, i.e. the term which is used within a given genus to dis-
tinguish any given species from every other species (or subspecies)
referred on taxonomic grounds to that genus.
This made the term "trivial name" a synonym of "specific epithet"
as used in bacteriology and botany. A radical change was directed at
the Copenhagen (1953) Zoological Congress which approved the
recommendation of the "Colloquium" that the expression "binomen"
be introduced, "defining that expression as denoting the binominal
combination which constitutes the scientific name of a species." In
other words, "binomen" was substituted for the previously approved
"specific name." The Copenhagen Congress further decided that
the "reversion to the use of the expression 'trivial name' was mis-
conceived." It repealed the Paris decision and substituted the ex-
pression "specific name" for "trivial name" on the basis that the
former expression had been released by the approval of the expres-
sion "binomen" for the name of a species as noted above. This action
means that in zoology "specific name" equates with "specific epithet"
in bacteriology and botany, and the zoological term "binomen"
equates with "specific name" or "species name" in other biological
sciences.

The use of an identical expression "specific name" for two dis-
tinct concepts, the one in zoology and the other in bacteriology and
botany, is unfortunate. Perhaps the difficulty lies in the zoological
use of the word "name." In the expression "trivial name" and the
later emendation "specific name" the word "name" is not intended
to designate a taxon.

The term specific epithet. This term, as used in bacteriology and
botany, requires examination and definition. The specific epithet is
the second component of the name of a species. Most often it is a
single word, but it may consist of two (or even more) words provid-
ing these words are together required to express a single concept or
description. A species name that has two or more unrelated words

Bacteria and Viruses 35

Rule 6. Sect. 1. Naming of Taxa

used as epithets is illegitimate; it is not a binary combination.
The species name having a single epithet made up of two or more
words as contrasted with a species name in which appear two or more
epithets may be illustrated. An organism belonging to the genus
Streptococcus that lives in sour milk is to be named. The two words,
sour milk, taken together name a single concept; together they con-
stitute a single epithet. From lac, Inctis (milk) and acidum (sour)
is derived an appropriate specific epithet, and the organism is named
Streptococcus lactis-acidi. The Streptococcus of lactic acid may be
termed Streptococcus acidi-lactici (from acidum lacticum = lactic
acid) . A common source of compound specific epithets in bacteriol-
ogy is formation from the names of diseases with which the organ-
isms may be associated. For the organism producing haemorrhagic
septicemia, Sternberg proposed the name Bacillus septicaerniae-haemor-
rhagicae. This name has a single specific epithet and does not con-
travene the directive of Rule 6 that a name of a species must be a
binary combination with only a single specific epithet. Pfeiler isolated
from the gangienous udder of a sheep a coccus which he named
Micrococcus mastitidis-gangraeyiosae-ovis, the Micrococcus of gang-
renous mastitis of sheep. Again, this is a single specific epithet, it is
the Latinized name of a disease, and the species name accords with
the rule.

The fact that a long compoinid specific epithet is acceptable in
the sense that it does not contravene a Rule does not mean that it is
desirable. Recommendation 6b (2) advises that specific epithets
should not be long. Long names are cumbersome. As a result of a
request from a gioup of bacteriologists working with microorganisms
associated with mastitis in cows, the shortened name Streptococcus
agalactiae Lehmann and Neumann was conserved against the older,
correctly formed binomial Streptococcus agalactiae-contagiosae Kitt.

A correct species name can have but one specific epithet; if more
than one is given the species name is illegitimate and unacceptable.
There may be confusion of three groups of species names:

(1) Legitimate species names in which the specific epithet is
made up of two or more words. In this case the two or more
words express a single idea and together modify the generic
name. Example: Pasteurella cholerae-gallinarum, the Pas-
teurella of cholera of fowls.

(2) Illegitimate species names in which there are two or more
specific epithets, the several epithets not related and sepa-
rately modifying the generic name. Example: Bacillus albus
lactis, the white Bacillus of milk. This cannot be made
legitimate by hyphenation of the two epithets to read Bacillus

36 International Code of Nomenclature

Rule 6. Sect. 1. Naming of Taxa

albus-Iactls, there are still two unrelated concepts. An at-
tempt to change the illegitimate trinomial to a legitimate
binomial yielded Bacillus nlbolactis. The name is a binomial
but now means the Bacillus of white milk, not the white
Bacillus of milk.
(3) Legitimate subspecies ?iames in which the generic name is
followed by a single specific epithet and this, in turn, by a
subspecific epithet, the whole ternary combination repre-
senting the name of a subspecies. This type of name will be
considered under Rule 7.
Hyphenation of compound specific epithets. Rule 6 states that
specific epithets made up of two or more words should either
be hyphenated or the words should be united. The Rule does not
indicate which choice should be made. Precedent indicates that
Avhen each component is separately declined, use of the hyphen is
advised. For example, if each component is in the genitive, as in
Clostridium oedematis-maligni , the hyphen indicates that the specific
epithet is made up of two separate words, the genitive of oedema
malignum, the name of the disease. Similarly one may write Strepto-
coccus lactis-acidi. Frequently one component modifies the other, as
an adjective preceding or following a noun, or is used in an adverbial
sense preceding a participle or an adjective, in such cases the two com-
ponents should be united into a single word, as nigromaculatus black-
spotted, thermophilus heat-loving, lentiputrescens slowly putrefying,
albogilvus whitish yellow. See Rule 27 and its Recommendations re-
lating to orthography.

Kinds of specific epithets. Rule 6 states that specific epithets may
be (1) adjectives, (2) nouns in apposition or (3) nouns in the geni-
tive.

(1) Adjectives as specific epithets. Specific epithets may be adjec-
tives, either simple or compound. They are of many origins, as
shown infra:

A simple Latin adjective: Staphylococcus aureus the golden
Staphylococcus; Synangium sessile the stalkless or sessile
Synangium; Neisseria sicca the dry Neisseria; Clostridium
botulinu7n the Clostridium related to sausage.
A compound Latin adjective: Sireptomyces alboflavus the whitish
yellow Sireptomyces; Bacillus flavoviridis the yellowish green
Bacillus; Rhabdomonas fusiformis the spindle-shaped Rhab-
domonas.
A simple Greek adjective (transliterated into Latin and with
Latin endings) Bacillus acinous the uncolored Bacillus; Bac-

Bacteria and Viruses 37

Rule 6. Sect. 1. Naming of Taxa

terium cliryseum the golden Bacterium: Microinotiospora
chalcea the bronze Micromoy\osporn.

A compound Greek adjective: Treponema calligyrum the beauti-
fully bent Treponejna; Streptomyces phaeochromogenes the
giay-color-producing Streptomyces: Pseudomonas cyanogenes
the dark-blue-producing Pseudomonas.

An adjective jormed jrom a place name: Brucella melitensis the
Maltese Brucella; Vibrio beroUnensis the Berlin Vibrio; Actino-
myces californiciis the Californian Actinomyces.

An adjective formed from the name of a person: Clostridium
pasteiirianiim the Pasteur Clostridium: Zoogloea beigeliana
the Beigel Zoogloea.

A participial adjective formed from a present participle: Eriuiyiia
citrimaculans the orange-spotting Envinia; Bacteroides co-
agulans the clotting Bacteroides.

A participial adjective formed from a past participle: Nocardia
maculata the spotted Nocardia: Clostridium malenominatum
the badly-named Clostridium.

(2) Substantives as specific epithets in apposition. A substantive
(noun) in the nominative case, in apposition with the generic name,
may be used as a specific epithet. This type of modifier is found in
most modern languages, as Victoria regina; London the novelist;
Burns the poet. In general the epithet expresses some characteristic
of the generic name with which it is in apposition; it has an adjectival
connotation. Examples: Bacillus radicicola. Bacillus the root dwel-
ler; Pseudomonas conjac the conjac Pseudomonas: Vibrio coynma the
comma Vibrio.

Some specific epithets taken from languages foreign to Latin are
of such form or spelling that they are not adaptable to Latin de-
clensions. Some words of this type were assimilated into Latin and
treated as undeclinable. For example, there are hundreds of per-
sonal and place names in the Latin Vulgate translation of the Bible.
The use of such undeclinable nouns as specific epithets is not in ac-
cord with Principle 4, which advises that as far as possible such names
be given appropriate Latin endings. In several genera of bacteria, as
in the genus Salmonella, it has become the custom to use the name of
the city or locality where the organism was first found as a specific
or subspecific epithet. There are hundreds of names stich as Salmo-
nella london, S. lexington, S. tennessee in the literature. For the
most part these organisms are differentiated on the basis of their
antigenic structure; they are infrasubspecific forms (serotypes) .

38 International Code of Nomenclature

Rule 6. Sect. 1. Naming of Taxa

The Enterobacteriaccac Subcommittee of the International Commit-
tee on Bacteriological Nomenclature is actively at work on the prob-
lems of taxonomy of these organisms. It seems probable that there
will finally be recognized a relatively small number of species, most
with many serotypes.

(3) Substa7itives in the genitive as specific epithets. The specific
epithet is often a noun in the genitive modifying the generic name.
It may be one of several kinds.

Genitive singular of a personal name: Borrelia kochii Koch's
Borrelia. If dedicated to more than one person of the same
name, the specific epithet may be in the plural. There are
several such instances in botany, but none has so far been
discovered in bacteriology.
Genitive singular of tJie name of a locality: Streptomyces novae-
caesareae the Streptomyces of Nova Caesarea (New Jersey) ;
Marmor angliae the Marmor of Anglia (England) .
Genitive of the name of a host, either plant or animal:

Genitive of the name of the genus of the plant host: Xantho-
monas antirrhini the Xanthomonas of Antirrhinum; Pseudo-
monas polygoni the Pseudomonas of Polygonum; Envinia
lathyri the Envinia of Lathyrus; Rhizobium phaseoli the
Rhizobium of Phaseolus.

Genitive of names of plant taxa i)i the plural: Rhizobium
leguminosarum the Rhizobium of the Leguminosae; Marynor
cruciferarum the Marmor of the Cruciferae; Xanthomonas
malvacearum the Xanthomonas of the Malvaceae.

Genitives of names of animals, singular: Corynebacterium bovis
the Corynebacterium of the ox; Haemobartonella canis the
Haemobartonella of the dog; Nocardia caprae the Nocardia
of the goat.

Genitives of the names of animals, plural: Arthromitus batra-
chorum the Arthromitus of frogs.

Genitives of names of diseases or lesions: Salmonella cholerae-
suis the Salmonella of cholera of the hog.

Genitives of other objects: Streptococcus cremoris the Strepto-
coccus of cream; Lactobacillus casei the Lactobacillus of
cheese; Pediococcus cerevisiae the Pediococcus of beer.

Illegitiynate vernacular genitives. In a few cases illegitimate ver-
nacular specific epithets have been introduced and have persisted for

Bacteria and Viruses 39

Rule 6. Sect. 1. Naming of Taxa

a time in the literature. For example, Duclaux (1882) described an
organism which he found associated with the production of slimy
or ropy milk. He placed it in his newly named genus Actinobacter
but did not give it a specific epithet. In consequence, the organism
has been termed Actinobacter du lait visqueux Duclaux, an illegiti-
mate combination.

40 International Code of Nomenclature

Recommendations 6a, 6b. Sect. 1. Naming of Taxa

RECOMMENDATION 6a. When it is desired to indicate the name of a sub-
genus in connection with the name of a species, the name of the subgenus
may be placed in parentheses between the two.

ANNOTATION

Recornini'iidatiou 6a. This reconmiendation is essentially that of
the Botanical Code and agrees with the corresjDonding rule in the
Zoological Code. The Bacteriological Code (like the Zoological)
recognizes the subgenus as the single category between genus and
species; the Botanical Code recognizes several intermediate categories,
for example, subgenus, section, subsection, etc. The corresponding
Recommendation of the Botanical Code states:

When it is desired to indicate the name of a subgenus or section
(or other subdivision to which a particular species belongs) in
connection with the generic name and specific epithet, the name of
the subdivision is placed in parentheses between the two (when
necessary, the rank of the subdivision is also indicated) .
Examples: Lactobacillus {Thermobacterium) caucasicus; Rickettsia
(Rickettsia) protoazekii; Rickettsia {Dermacentroxe7ius) rickettsii.

RECOMMENDATION 6b. In forming specific epithets bacteriologists should
attend to the following recommendations:

(1) To choose a specific epithet which, in general, gives some indication
of the appearance, the characters, the origin, the history, or the
properties of the species. If taken from the name of a person, it
usually recalls the name of the one who discovered or described it
or was in some way concerned with it.

(2) To avoid those which are very long and difficult to pronounce.

(3) To avoid those which express a character common to all or nearly
all the species of a genus.

(4/ To avoid using the names of little-known or very restricted localities,

unless the species is quite local.
(5) To avoid, in the same genus, epithets which are very much alike,

especially those which differ only in their last letters.
(6} Not to adopt unpublished names found in authors' notes, attributing

them to their authors, unless these have approved publication.
(7) To avoid compound specific epithets which include word stems from

two or more languages (epitheta hybrida).

Bacteria and Viruses 47

Recommendation 6b. Sect. 1. Naming of Taxa

ANNOTATIONS

Recommendation 6b (1) . Examples: Staphylococcus aureus, Clo-
stridium pasteurianum, Xanthomouas campestris. Bacillus rnsccjsus,
Kiirthia zopfii.

Recommendatioyi 6b (2) . Notwithstanding pronunciation difficulties
and use of consonant combinations quite foreign to classic Latin, we
accept a name such as Cytophaga krzemieJiiewskae. Madam Krze-
mieniewska was a noted contributor to our knowledge of certain
groups of microorganisms, particularly the myxobacters.

Some specific epithets derived from names of diseases are long
and cumbersome, as in the species names Bacillus pneumoenteritidis-
miirium (Bacillus of pneumoenteritis of mice) , Bacillus septicaemiae-
anserum-exudativae (Bacillus of exudative septicaemia of geese) , or
Leptospira icterohaemoglobinuriae (Leptospira of icterohaemoglobi-
nuria) . Two of these specific epithets have ten syllables and one has
fourteen. Many long names were not validly published, or are for
some other reason illegitimate.

Recommendation 6b (3) . Rarely has this suggestion been ignored.
A name such as Micrococcus sphaericus is obviously not distinctively
descriptive of the species since presumably all species of this genus
have spherical cells.

Recomynendation 6b (4) . This recommendation, taken from the
Botanical Code originally, has relatively little significance in bac-
teriology. One may not know that the Vineland of Azotobacter vine-
landii is in New Jersey. In some cases authors have chosen to use
the mediaeval or late Latin place names, making the modern meaning
obscure.

Recojnmendation 6b (5) . Confusion of specific epithets in the same
genus due to similarity in spelling has not been common in bacteri-
ology. A hypothetical case may illustrate. There is a named bac-
terial species Pseudoynonas barkeri, likewise a genus of orchids
named Barkeria. Suppose a new species of a pseudomonad parasitic
upon Barkeria were named Pseudomonas barkeriae. There might
well be some confusion with two specific epithets in the same genus
so closely resembling each other as barkeri and barkeriae. Even
more difficult to differentiate in pronunciation would be the recog-
nized Pseudomonas woodsii and a hypothetical P. ivoodsiae named
from the fern genus Woodsia.

42 International Code of Nomenclature

Recommendation 6b. Sect. 1. Naming of Taxa

Reconiinendation 6b (6) . This recommendation, found necessary in
botany, has rarely, if ever, been of significance in bacteriology.

Rccommcndatioyi 6b (7) . Many such hybrid words used as specific
epithets have been proposed, as acidophilus (acid-loving) , nlbo-
chryseus (whitish golden) .

Additional Recommendations of the Botanical Code. The Botanical
Code includes several Recommendations relating to specific epithets
that are of interest. The Botanist is urged:

(a) To use Latin temiinations insofar as possible.

(d) To avoid those formed of two or more hypliened words.

(e) To avoid those which have the same meaning as die generic
name (pleonasm) .

(h) To avoid those which haAe been used before in any closely allied
genus.

Additional Recommendations from the Zoological Rules are also
pertinent.

(1) The best specific name (epithet) is a Latin adjective, short,
euphonic, and of easy pronunciation. Latinized Greek words
or barbarous words may, however, be used. Examples: gym-
nocephalus, echinococciis, agnti, koactli, urubitinga.

(2) It is well to avoid the introduction of the names (epithets)
typicus and typus as new names for species or subspecies,
since these names are always liable to result in later con-
fusion.

(3) The use of compound proper names indicating dedication,
or of compound words indicating a comparison with a simple
object, does not form an exception to Art. 2. In these cases
the two words composing the specific name (epithet) are
written as one word with or without the hypen. Examples:
sanctae-catharinae or sanctaecatliarinae , jan-maveni or
janmaveni, cornu-pastoris or cornupastoris, com-anguinum or
cornanguiniim, cedo-nulli or cedonulli.

(4) Expressions like rudis planiisque are not admissible as spe-
cific names (epithets) .

Bacteria and Viruses 43

Recommendation 6c. Sect. 1. Naming of Taxa

RECOMMENDATION 6c. Avoid the use of the genitive and the adjectival
form of the same specific epithet to designate two different species of the
same genus.

ANNOTATIONS

Recommendation 6c. A specific epithet derived from the name of a
person or locahty may be a noun in the genitive or it may be con-
verted into an adjective by use of an appropriate ending. There is
a Bacillus pasteurii and a Clostridium pasteurianum. Should either
species be transferred to the other genus there would be some danger
of confusion. The two specific epithets beigelii and beigelianum have
been used for the same species in combination with different generic
names. In bacteriology a specific epithet is rarely in adjectival form
when derived from the name of a person. Conversely, specific epithets
from place names are rarely in the genitive.

44 International Code of Nomenclature

Rule 7. Chapter 3. Sect. 1. Naming of Taxa

RULE 7. Names of subspecies (varieties) are ternary combinations consisting
of the name of the genus followed by specific and subspecific epithets in
order.

Epithets of subspecies (varieties) are formed like those of species; when
adjectival in form and not used as substantives they agree in gender with
the generic name.

Neither within the same species nor within the same genus may two
subspecies bear the same subspecific epithet.

If the species is divided into subspecies, the subspecific epithet of the
subspecies containing the type of the species must be the same as that of
the species.

NOTE. Names of species and of subspecies (varieties) from a nomenclaturol
point of view are coordinate (of equal value) and are subject to the same
rules and recommendations.

NOTE. The words subspecies and variety are alternative designations for
the same taxonomic rank; they are synonyms. The use of the term sub-
species is preferable.

ANNOTATIONS
Rule 7. The name of a subspecies (or variety) is a trinomial made
up of a generic name followed by a specific epithet and this, in turn,
by a subspecific epithet. As provided in Rule 14, an author when
proposing the name of a subspecies should make clear its subspecific
status. The epithets preferably are separated by the abbreviation
indicating a subspecies (variety) , or this may be omitted. One may
write Bacillus subtilis subsp. 7nger or Bacilhis subtiUs var. niger or
Bacillus subtilis niger. The last listed method is most often used in
biology, but has the disadvantage of not being directly distinguish-
able from an illegitimate trinomial name of a species. In other words,
the name Bacillus fluorescens liquefaciens proposed as the name of a
species was illegitimate because it had two specific epithets; it was
not proposed as the name of a subspecies.

The sixth edition of Bergey's Manual of Determinative Bac-
teriology recognized two varieties or subspecies of Bacillus subtilis:
Bacillus subtilis aterrimus and B. subtilis Jiiger. A third should have
been recognized, B. subtilis subtilis, to inckide the type of the species.

Note that species may be divided into subdivisions of lower rank
than subspecies and that subspecies may also be subdivided. These
subdivisions are considered under Rule 8. Not all of the rules and
recommendations applicable to species and subspecies are applicable
to taxonomic ranks lower than subspecies.

Recommendations 6a, 6b and 6c apply to subspecific epithets as
well as to specific epithets.

The name of a subspecies is a ternary combination; the use of a
binary combination as the name of a subspecies is therefore not ad-
missible. Example: Mycobacterium tuberculosis var. avium or Myco-
bacterium tuberculosis avium, not Mycobacterium tuberculosis var.
M. avium,.

Bacteria and Viruses 45

Chapter 3. Sect. 1. Naming of Taxa

RULE 8. Infrasubspecific subdivisions. Subdivisions of species (other than
subspecies [varieties]) and subdivisions of subspecies are given vernacular
names or designated by numerals or letters or, in special cases, are given
names in Latin form. These are termed infrasubspecific subdivisions or forms.
The names given to organisms included in infrasubspecific subdivisions need
not conform to the rules governing the naming of subspecies and higher
taxa. These rules do not determine the naming of forms of infrasubspecific
rank.

NOTE. These subdivisions are regarded as of lower rank than subspecies.
The terms commonly used to designate them are strain, biotype, serotype,
morphotype, phagotype, group, phase, forma (or forma specialis), variant,
mutant, stage, state.

ANNOTATIONS

Rule 8. The rules and recommendations of the three biological
codes of nomenclature relating to subdivisions of species and sub-
species are not uniform.

Bacteriological Code. The category subspecies is the lowest sub-
division of species recognized and governecl by all the Rules and
Recommendations of this Code; "variety" is regarded as a synonym
of "subspecies." Various terms indicated in the Note to Rule 8 above
are of lower than subspecific rank. A collective name has not been
recognized for the designation of subdivisions of lower rank than
subspecies. The names given to infrasubspecific forms are frequently
not Latin and are not subject to the rules governing the naming of
bacteriological taxa. The term "infrasubspecific" was proposed in the
Zoological Rules to include these lower subdivisions and is appropri-
ate for use in bacteriology as well. Confusion will be avoided if
bacteriologists will use the names only in a generally accepted sense,
as indicated in Recommendation 8a.

Botanical Code. The botanists recognize a whole hierarchy of
names of taxa subordinate to species. These are subspecies, variety,
subvariety, forma, subjorma. An example given is: "Saxifraga aizoon
var. aizoon subvar. brevi folia forma multicauUs subforma surculosa
Engler et Irmscher."

The Botanical Code states that the use of a binary combination
for an infraspecific taxon is not admissible, but ternary combinations
may be used provided the rank of the taxon is stated. For example,
using the illustration given above, "Saxifraga surculosa Engler et
Irmscher" is inadmissible, but "Saxifraga aizoon subforma surculosa
Engler et Irmscher" is in proper form.

46 International Code of Nomenclature

Rule 8. Sect. 1. Naming of Taxa

The Botanical Code (Art. 37) has the following pertinent statement:
Two subdivisions of the same species, even if they are of different
rank, cannot bear the same subdivisional epithet, unless they are
based on the same type. If the earlier subdivisional name (ternary
combination) was validly published, the later one is illegitimate
and must be rejected.

Zoological Rules. The Zoological Code includes all taxonomic
ranks lower than subspecies under the general term infrasiibspecific.
An infrasubspecific form is defined as any subdivision of a species
other than a subspecies, such as a "seasonal, sexual, or transitional
form, an aberration, or oj;her minority element within a species."
The Zoological Code also includes a number of Rules and Recom-
mendations relating to the use and recognition of infrasubspecific
names (epithets) .

(1) Ti7ne of publication. The recognition of a distinct infrasub-
specific category of names is recent. The problem is faced as to
the allocation of names (epithets) as between subspecific and
infrasubspecific. Names given before 1951 and after 1950 are
treated separately.

Publication before 1951. A name is to be regarded as that of
a subspecies if the author so indicated or if he did not indi-
cate whether he regarded it as a subspecies or an "infrasub-
specific" form. It is to be recognized as infrasubspecific only
if expressly so indicated. The ultimate test is whether the
name (epithet) was applied to a population (subspecific) or
to some minoritv element within the population (infrasub-
specific) .

Publication after 1950. Names of less than specific rank, in-
cluding those proposed as varieties, are to be regarded as in-
frasubspecific unless definitely designated as the names of
subspecies.

(2) Nomenclatural status.

a. Names of species and subspecies on the one hand and the names
of infrasubspecific forms on the other constitute mutually inde-
pendent sections of nomenclature as regards both the Law of
Priority and the Law of Homonymy.

b. A name of an infrasubspecific fomi is to be coordinate with the
names of all other infrasubspecific forms, but not with the names
of subspecies or species.

c. A name originally published as that of an infrasubspecific form
may be elevated by a svibsequent reviser to the status of a sub-
specific name or a specific name. In that event it is to rank (in
its new status) , for purposes of priority, from the date on which
it was so elevated, and it is to be attributed to the author by
whom it was so elevated.

Bacteria and Viruses 47

Rule 8. Sect. 1. Naming of Taxo

There are also included several Rules and Recommendations
relative to the form of acceptable publication and of citation of
infrasubspecific names. The Rules also provide a formula for the
citation of the name of an infrasubspecific form.

A citation of the name of an infrasubspecific form is to include
(a) its binomen (if it is treated as a form of a subspecies, its tri-
nomen) , (b) after the specific (or subspecific) name, a comma fol-
lowed by an expression of the status attributed to the form in
question, and (c) the name of the inhasubspecific form.

48 International Code of Nomenclature

Recommendation 8a. Sect. 1. Naming of Taxa

RECOMMENDATION 8a. Authors of names of subdivisions of species of
bacteria which are not treated as subspecies (varieties) should attend to the
following recommendations and defmitions:

(1) A strain is made up of the descendants of a single isolation in pure
culture. It may be designated in any manner, as by the name of the indi-
vidual responsible for its isolation (as Corynebacterium diphtheriae strain
Park-Williams); by the name of a locality, by a number, or by some similar
laboratory distinguishing mark. "Strain" may also be used to designate cul-
tures of bacteria which correspond to cultivated "varieties" (cultivars) of
higher plants in having some special economic significance.

ANNOTATIONS

Recnmrnendntion 8a (1) . Rule 9 emphasizes the importance of pre-
serving the particular strain of a species which was first isolated and
studied in the preparation of the species description, i.e., the type
culture of the species.

A particular strain of a species or subspecies may show marked
differences in economic significance from other strains or isolations.
Corynebacterium diphtheriae strain Park-Williams has long been
used for commercial production of diphtheria toxin. The yield of
the antibiotic streptomycin varies greatly with the strain of Strepto-
myces griseiis used in its production. Strains of organisms which
show marked differences in virulence, pigment production, enzyme
yield, host specificity, host preference, or other characters may be
labeled appropriately.

Certain mutant strains of microorganisms may require special
identification because of their importance in providing a biochemical
marker for cell-metabolism studies. For example, by irradiation of
a bacterial culture followed by appropriate techniques of isolation,
it may be possible to secure a mutant strain that differs from the
parent culture in inability to synthesize leucine and thus exhibits a
growth requirement for this amino acid in the cultiue medium. An
organism having this characteristic may be known as "leucineless
strain."

A strain which originates as a mutant may also be termed a
variant, though the term variant does not necessarily indicate a true
gene mutation. Variants are constantly arising, such as differently pig-
mented sectors in a colony, or a lactose-fermenting mutant in a culture
of a non-lactose-fermenting organism.

Bacteria and Viruses 49

Recommendation 8a. Sect. 1. Naming of Taxo

RECOMMENDATION 8a (2). Type is a term which has frequently been used
to designate a subdivision of a soecies, particularly in cases where the
distinguishing characters are regarded as insufficient to justify the recog-
nition of a subspecies. Types are often distinguished on the basis of anti-
genic characters. However, in view of the preferred use of the word "type"
in a different sense as defined in Principle 1 1 and Rule 9, it is recommended
that the terms serotype (or serological type), biotype (or physiological type),
phagotype (or phage type) and morphotype (or morphological type) be ap-
propriately substituted for type as a designation of a subdivision of a species.

ANNOTATIONS

Recojnmendation 8a (2) . The continued use of the word "type" in
microbiology with two very different meanings is a major source of
confusion. The preferable and recommended use of "type" is that
in the sense of "nomenclatural type" of a taxon and in phrases such
as "type culture" and "type species", as noted in Principle 11 and par-
ticularly Rule 9. This use of the word "type" is recommended.

The second common and less desirable use of the word "type"
is for the designation of certain strains or cultures or groups of these
within a species which show certain distinctive characters of less than
subspecific rank. Certain substitute terms are recommended.

A "serotype" is a subdivision (infrasubspecific) of a species or
subspecies distinguishable from other strains of the same species on
the basis of its antigenic structure. It may be suitably named by
use of the species name followed by '"serotype" and this in turn bv
a suitable epithet or formula, as, for example. SItigella flexneri sero-
type la.

A "biotype" is a subdivision of a species, subspecies or serotype
which may be distinguished from other subdivisions of the species
by possession of some special or usefully diagnostic physiological
character. An anaerogenic mutant of an aerogenic species of bacteria
might be named by following the species name by "biotype anaero-
genes" (or, anaerogenic biotype) .

A "morphotype" is a subdivision of a species distinguished by
possession of some special or unusual morphological character which
may or may not be associated with a change in serological state. A
capsulated variant of a bacterial species in which the cells are
normally noncapsulated might be named by use of the species name
followed by "'morphotype capsiilatus."

A "phagotype" is a subdivision of a species which is distinguished
by its sensitivity to a particular bacteriophage or by a distinctive
pattern of sensitivity to a set of specific bacteriophages. It may be
characterized by the carriage of a symbiotic phage.

50 Infernational Code of Nomenclature

Recommendation 8a. Sect. 1. Naming of Taxa

RECOMMENDATION 8a (3). The term "group" in bacteriology should be
used with great care and be well defined if ambiguity is to be avoided. It has
been used in somewhat different senses by those working in various fields
of bacteriology. "Group" is used to designate congeries of organisms having
common characteristics. In many cases the groups are based upon antigenic
analyses, they are assemblages of related serotypes.

ANNOTATIONS

Recommendation Sa (3) . Among the several bacteriological uses of
"group" are the following:

(1) "Group" is used by the Enterobacteriaceae Subcommittee of
the International Committee on Bacteriological Nomenclature
to emphasize the belief that recognition and diagnosis of the
many serotypes is of fundamental significance. Those having
some similarities in antigenic structure are placed together
in "subgroups," then by use of various physiological tests
larger groups are formed. The committee uses such designa-
tions as Salmonella group, Escherichia group, Shigella group
where other bacteriologists would use the designations genus
Salmonella, genus Escherichia, etc. An effort is made to avoid
in some cases the conventional terminology of biology for
what are regarded as good reasons.

(2) Workers with certain bacterial genera and families have used
the term "group" in antigenic analysis to designate species or
subgenera (e.g., Streptococcus group A Lancefield) , or varie-
ties or subspecies (e.g., Neisseria meningitidis group I Scott) .
It has been suggested that the term gi^oup be reserved for
primary serological divisions and designated by capital let-
ters. Any serological subdivisions within the group should
be designated as serotypes and distinguished by Arabic num-
bers (e.g., Streptococcus pyogenes Rosenbach group A Lance-
field, serotype 1 Griffith) .

The term "group" has not as yet achieved in bacteriology a gen-
erally accepted and consistent definition.

Bacteria and Viruses 51

Recommendation 8a. Sect. 1. Naming of Taxa

RECOMMENDATION 8a (4). The designation phase should be restricted to
well-defined stages of a naturally occurring alternating variation.

ANOTATIONS

Recommendation 8a (4) . Andrewes described diphasic \ariation of
the flagella of many serotypes of SalmoneUa. This form of variation
appears to be limited to the H (flagellar antigens) of species of
Salmonella and certain closely related bacteria (Arizona group
[Edwards]) . The use of the term "phase" for a more permanent vari-
ation (e.g., S — ^R, V — ^W, "phases" of Haemophilus pertussis) is un-
desirable. Phase Aariation as here defined is thus far unknown outside
the family Enterobacteriaceae.

RECOMMENDATION 8a (5). A form (forma) or special form (forma specialis)
is a subdivision of a species of a parasitic or symbiotic microorganism dis-
tinguished primarily by adaptation to a particular host. It is named prefer-
ably from the scientific name of the host written in the genitive.

ANNOTATIONS

Recommendation 8a (5) . This recommendation has proved useful
in the naming of certain fungi which are selective as to host plant,
but which show little or no consistent morphological differences.
Some plant rusts, for example, are to be found on a number of
species of grasses or \arieties of the grasses, but they are physiologic-
ally so specialized or adapted that the forma found on one species or
variety of host will not grow on some other. No adequate morpho-
logical distinguishing character is known. A form so specialized as
to pathogenicity may be termed a forma specialis, designated by
"f. sp.," followed by the name of the host species to which it is
adapted. This method of identification has only occasionally been
used in naming bacteria, though it may be useful in distinguishing
bacterial plant parasites or symbionts. For example. Fang, Allen,
Riker and Dickson noted a forma specialis, Xanthomonas translucens
f. sp. phlei-pratensis on the grass timothy {Phleum pratense) .

RECOMMENDATION 8a (6). State (or stage) is the name given to the
rough, smooth, mucoid and similar variants which arise in cultures of many
species of bacteria. These are usually regarded as alternating states which
are often reversible, and indeed by some authors considered as part
of a pleomorphic life cycle. They may be designated by some vernacular
descriptive name.

Section 2

Designation of Nomendatural Types

RULE 9a. For each taxonomic group (taxon) there shall be designated a
ncmenclatural type. The nomendatural type of a class or a subclass is an
order, that of an order or suborder is a family, that of a family, subfamily,
tribe or subtribe is a genus, that of a genus or a subgenus is a species, that
of a species or subspecies is preferably an authentic culture, but it may be
a specimen or preparation, illustration or description.

ANNOTATIONS

Rule 9a. The type concept has increasingly become the basis for
all modern biological taxonomy. When taxa of any rank are divided
or united, the use of the nomendatural type concept leads to rational
determination of the correct names to apply. Increasingly, stability
and rationality in microbiological nomenclature is dependent upon
maintenance of adequate type culture collections.

The directive of Principle 11, which states that for each named
taxon there shall be fixed a nomendatural type, is applied in Rule
9. The definition given in Principle 11 is:

A nomendatural type is that constituent element of a taxon to

which the name of taxon is peimanently attached.
The meaning of the phrase "permanently attached" may be illus-
trated. The generic name Bacillus is the nomendatural type of the
family BaciUaceae. The family name Bacillaceae is "permanently
attached" to the name of the genus Bacillus in the sense that there
can be no family Bacillaceae that does not include the genus Bacil-
lus. As usually interpreted the family Bacillaceae may include not
only the genus Bacillus but also such other genera as are regarded
as being sufficiently closely related in their characteristics to the
genus Bacillus. The placement of Bacillus in Bacillaceae is objective,
while the inclusion of any other genus is subjective; it depends on
the decision of the student as to whether the other genus resembles
Bacillus sufficiently. The genus Bacillus is the permanent element
within the family Bacillaceae.

Similar is the meaning of "nomendatural type" as applied to the
type spedes of a genus. In the genus Bacillus the type species is
Bacillus suhtiUs, and the generic name Bacillus is "permanently at-
tached" to the species Bacillus subtilis. The genus Bacillus may be
defined as the species Bacillus subtilis together with such other species
as are recognized as having certain characters in common with those
of Bacillus subtilis.

RULE 9b. Designation of the nomenclotural type of a taxon of higher rank
than genus. The nomendatural type of a family, subfamily, tribe or sub-
tribe, whose name has been formed in conformity with the Rules, is that
genus from whose name the name of the higher taxon has been derived.
The nomen^-iatural type genus of a family whose name has been conserved,

[53]

54 International Code of Nomenclature

Rules 9b, 9c. Designation of Nomenclatural Types

but was not formed in conformity with Rule 3, may be fixed by the Judicial
Commission. The nomenclatural type of a taxon of rank higher than family
may be fixed at the time of its proposal by the author. If not so fixed, it
may be designated by the Judicial Commission.

NOTE. If subdivisions of families (subfamilies, tribes, subtribes) are recog-
nized, one taxon of each category must include the type genus of the
family and its name should be derived from the name of the type genus.

ANNOTATIONS

Rule 9b. The family name Bacilloceae is derived from the generic
name Bacillus, and this is the name of its nomenclatural type. There
may be a subfamily BaciUoideae, a tribe Bacilleae and a subtribe Bacil-
linae, all with the nomenclatural type Bacillus.

The nomenclatural type of none of the named subclasses, orders
and suborders in bacteriology has thus far been definitely fixed either
by designation or by action of the Judicial Commission.

The family name Enterobacteriaceae has been placed on the list
of genera consemayicla to replace the rejected family name Bac-
teriaceae.

RULE 9c. Designation of the nomenclatural type of a genus or subgenus.

(1) The nomenclatural type (type species) of a genus or subgenus is the
name of the single species or of one of the species included when
the name of the genus or subgenus was originally validly published.
NOTE. The expression "type species" is to be used rather than
"genotype" or other expressions when referring to the type species
of a genus.

(2) If the author in the original publication of a generic or subgeneric
name definitely selected a type species, this species shall be accepted
as the nomenclatural type (type species) regardless of other con-
siderations (type selection by original designation).

NOTE. The meaning of the expression "selected a type species" is

to be rigidly construed. Mention of a species as an illustration or

example of a genus does not constitute selection of a type.

(3) If the author of a generic or subgeneric name in his original publica-
tion failed to designate a type species, the type may be selected by
a subsequent author, and the author who first makes the choice
must be followed unless it can be proved that his choice is not in
accordance with the following rules:

(a) If the genus, when originally published, included but one species,
this species shall be the nomenclatural type (type species) (selec-
tion by monotypy).

(b) if the genus, when originally published, included more than one
species, the type species selected shall be one of these. However,
species inquirendae, species doubtfully referred to the genus,
species mentioned as in any way exceptional, species which
definitely disagree with the generic description (provided others
agree), and species which possess characters stated in the generic
description as rare or unusual are to be excluded from considera-
tion in selecting the type.

Bacteria and Viruses 55

Rule 9c. Designation of Nomenclatural Types

(c) If none of the species named by an author in his original de-
scription and publication of a generic name can be recognized,
that is, if no identifiable type species can be selected in accord-
ance with the rules, the Judicial Commission may issue an Opinion
declaring such generic name to be a nomen rejiciendum and
without standing in nomenclature. However, if such generic
name has come into use for identifiable species named subse-
quently, one of these later species may be selected by inter-
national agreement as the type species with the generic name
ascribed to the author of the binomial designated as the type
species. Such selection of a type species and recognition of the
author of the generic name must be based upon an Opinion by
the Judicial Commission, and if such Opinion is questioned, its
validity shall be determined by action of the International Com-
mittee on Bacteriological Nomenclature.

(d) The publication of a new generic name as an avowed substitute
for an earlier one does not change the type of the genus.

ANNOTATIONS

Rule 9c. The use of the type method as a guide in taxonomy and
nomenclature was first adopted by the zoologists, then by the botan-
ists. Recognition of the principle by the bacteriologists came later, in
part because of the difficulty in determining what should constitute
the type of a species. Obviously no nomenclatural types were desig-
nated for the genera described by the early ^vorkers in bacteriolog) .

Botanical rules relating to the designation of type species. The
Botanical Code and the Bacteriological Code are substantially the
same in essentials. The Botanical Code provides that if the author of
the generic name did not designate a type species, but gave to a spe-
cies a specific epithet typicns or typus, that species shall be regarded as
the type species.

Zoological rules relating to designation of type species. While
quite differently phrased and much more complex and detailed, the
Zoological Rules do not differ markedly in essentials from those of
the Bacteriological Code.

The Zoological Rules permit tautonomy, that is, the use of the
same word for the generic name and for the specific name (specific
epithet) of the name of a species, as in the name Trutta trutta.
Tautonomy is forbidden in Botany and Bacteriology (Ride 25) .

The Zoological Rules state:

If a nominal genus, witliout originally designated or indicated type,
contains among its original nominal species one possessing the
generic name as its specific or subspecific name, either as its oldest
available name or as a synonym, that normal species or subspecies
becomes ipso facto type of the nominal genus.

The Zoological Rules also include a Recommendation with a
list of precepts that should be followed in order of precedence in

56 International Code of Nomenclature

Rules 9c, 9d. Designation of Nomenclaturol Types

designation of a type species by subsequent selection. Few or none

of these seem to be applicable in bacteriological nomenclature.

A rule in zoology definitely fixes the use of the designation "type

species." It reads:

The concept of a type species of a genus is invariably to be indi-
cated by the expression "type species."

The following directive is also included as a Recommendation:
When referring to the type species of a genus, a zoologist should
always use the expression "type species" rather than "genotype"
or any other expression.

RULE 9cl. The nomenciatural type of a species or subspecies. The nomen-
clatural type of a species or subspecies may be a living culture maintained
in a bacteriological laboratory, more particularly in one of the international
or national type culture collections.

NOTE. For a species which cannot be maintained in laboratory cultures or
for which neither type cultures nor neotype cultures exist, the type is the
original description, preparation or illustration.

NOTE. Definitions.

(a) The term cuhure is to be interpreted as including every method of
maintaining organisms in a living state (in a medium, in a host by
passage, in cells or exudates, or desiccated).

(b) A type cuhure is a living culture of an organism which is a descendant
of the original culture or an isolation from which the author who first
described the organism made his original description, which culture
has been maintained pure, and which agrees in its characteis with
the original description.

(c) A neotype cuhure is one which has been accepted by international
agreement to teplace a type culture which is no longer in existence.
It should agree with the diagnosis given by the original describer
and should be recommended by those workers familiar with the
species, and their agreed recommendation approved by the Judicial
Commission.

ANNOTATIONS

Rule 9d. Botanists and zoologists have de\eloped a series of names

to be used in connection with the fixing of type specimens of species

of plants and animals.

Holotype (From the Greek, loholc or entire, the getiiiive type) .

This is defined by the Botanical Code as follows:

A holotype (type) is the one specimen or other element used by
the author or designated by him as the nomenciatural type. For so
long as a holotype is extant it automatically fixes the application of
the name concerned.

I he definition given in zoology is:

The single specimen designated or indicated as "the type" by the
original author at the time of the original publication.

Bacteria and Viruses 57

Rule 9d. Designation of Nomencloturol Types

The term holot^pe has found little use in bacteriology inasmuch
as the nomenclatural types of bacterial species obviously cannot be
the dried or preserved specimens used for higher plants and animals.
A bacterial type should, if possible, be a living culture. This re-
quirement complicates the picture of type designation in bacteriology
and emphasizes the importance of Type Culture Collections being
what the name indicates, collections containing authentic type cul-
tures.

The definition of "type culture" given above in Rule 9 may be
regarded as the bacteriological equivalent of the definition of a holo-
type.

Lectotype (From the Greek, chosen or selected) . The Botanical
Code reads:

A lectotype is a specimen or other element selected from the original
material to serve as nomenclatural type when the holotype was not
designated at the time of publication or for so long as it is missing.
The Zoological Rules define lectotype as:

A single specimen, selected, after the original publication, from a
series of syntypes to be "the type"; such selection, in order to be
effective, to be made known by publication.
The term lectotype has found little or no use in bacteriology.
Neotype (From Greek, new) . The Botanical Code states:
A neotype is a specimen selected to serve as nomenclatural type for
so long as all of the material on which the name of the taxon was
based is missing.
The zoological definition is:

Neotype: A single specimen, identified with a nominal species al-
ready described and designated under the prescribed procedure
as a unique standard of reference to replace a holotype or lectotype
believed to be lost or destroyed.

.\ large proportion of the type cultures of the species of bacteria
will necessarily be neotypes; authentic descendent cultures from origi-
nal isolations by authors of names are frequently unavailable. Un-
fortunately, few of the cultures maintained in the recognized Type
Culture Collections are authentic type cultures. In relatively few
cases have type cultures been definitely designated. Study and defi-
nite designation of the type cultures (largely neotypes) is a necessary
step in stabilization of bacteriological nomenclature.
Syntype is defined in zoology as:

One of a number of specimens of equal nomenclatural rank which
formed all or a part of the material before the original author, in
those cases where that author did not designate or indicate a holo-
type.
In botany the statement is:

One of two or more specimens or elements used by the author
when no holotype was designated, or one of the two or more speci-
mens simultaneously designated as type.

58 International Code of Nomenclature

Recommendations 9a, 9b. Designation of Nomenclatural Types

RECOMMENDATION 9a. When publishing the name of a new taxonomic
group (taxon), authors should designate carefully the nomenclatural type
of the taxon being named. This type determines the application of the name
in the event of this taxonomic group being subsequently divided.

ANNOTATIONS

Recommendation 9a. In recent years many authors of new generic
names have designated the type species. Castellani and Chalmers
(1919) definitely designated Bacterhim coli commime Escherich as
the nomenclatural type of their new genus Escherichia with the name
Escherichia coli (Migula) Castellani and Chalmers.

Waksman and Henrici named a new genus Streptomyces, with the
type species Streptomyces albiis (Rossi-Doria) Waksman and Hen-
rici (Streptothrix alba Rossi-Doria, Actinomyces albns (Rossi-Doria)
Krainsky) .

RECOMMENDATION 9b. The utmost importance should be given to the
preservation of the original "type" material on which the descriptions of
new species and subspecies are based. If the microorganism is one which
may be maintained in pure culture, an authentic culture designated as the
type culture should be deposited with one or more of the national or inter-
national type culture collections.

Bearing in mind the morphological, biochemical, antigenic and virulence
changes that may occur as the result of repeated subculture, every pre-
caution should be taken to maintain such cultures with a minimum amount
of change. It is likewise important that descriptions, illustrations and diag-
noses of new species and subspecies be as complete as possible.

ANNOTATIONS

Recommendation 9b. Corresponding statements in the Botanical

Code read:

It cannot be too strongly recommended that the original material,
especially the holotype, of a taxon be deposited in a permanent re-
sponsible institution and that it be scrupulously preserved. Where
living material has been designated as a type, appropriate parts of
it should be immediately preserved.
The Zoological Rules have several provisions relative to type

specimens.

Holotypes, syntypes and lectotypes are the property of science.
Upon publishing a description of a new species, subspecies or "in-
frasubspecific" form, an author should affix a conspicuous label to
the holotype or lectotype, indicating its type status, and should
deposit the specimen in a museum or other institution where it
will be safely preserved and will be available for study.

Publication of Names

RULE 10. Valid publication is treated as beginning for the bacteria with 1
May 1753, the date of publication of Linnaeus' Species Plantorum, edition 1.

ANNOTATIONS

Rule 10. The problem of fixing the beginning date for valid publi-
cation of names for plants was considered at several of the early
Botanical Congresses. It was agreed that names published before the
date of Linnaeus' Species Plantariim 1753 would not be considered
as validly published. The two volumes of the Species Plantariim
appeared respectively in May and August of 1753. For nomenclatural
purposes it was decided that both volumes should be regarded as
having been published simultaneously on 1 May 1753.

However, it was evident that the Species Plantariim did not give
an adequate coverage of certain groups of plants, e.g., fungi and algae.
It was decided to accept certain monographic treatments published
at later dates as the beginnings of valid publication for such groups.
As fixed in the Botanical Code, valid publication of the groups of
interest to microbiologists are as follows:

Fungi: Uredinales, Ustilaginales and Gasteromycetes, 31 Dec. 1801.

(Persoon, Synopsis Methodica Fungorum) .
Fungi Caeteri, 1 Jan. 1821 (Fries, Systema Mycologicum, Vol. 1) .
Algae: Nostocaceae Homocysteae, 1892-93 (Gomont, Monographic

des Oscillaries, Ann. Sci. Nat. Bot. VI. i5:263-368; 76:91-264) .

Nostocaceae Heterocysteae, 1886-88 (Bornet and Flahault, Re-
vision des Nostocacees heterocystees. Ann. Sci. Nat. Bot. VII.

3:323-381; ^:343-373; 5:51-129; 7:177-262).

Desmidiaceae, 1848 (Ralfs, British Desmidieae) .

Oedogoniaceae, 1900 (Hirn, Monographic und Iconographic dcr

Oedogoniaccen. Acta Soc. Sci. Fenn. 27 [1]).

Myxomycetes, 1 May 1753 (Linnaeus, Species Plantarum ed. 1) .

Bacteria, 1 May 1753 (Linnaeus, Species Plantarum, cd. 1) .
At the Botanical Congress held in Brussels in 1910, various "points
of departure" for the bacteria were considered. Vuillemin proposed
that 1753 be adopted. Klebahn suggested that the work of F. Cohn
(1870-76, Untersuchungen iiber Bakterien) was more suitable. His
position was approved by Engler. Magnus contended that much work
of systematic value had been accomplished before 1870 and proposed
that the publication of Ehrenberg in 1786 [1838] should be the
starting point. (Possibly Magnus intended Mueller 1786.) Vuillemin
contended that if a date subseqtient to 1753 be taken, it might well
be 1910. The whole matter was finally deferred to the next Congress.

[59]

60 International Code of Nomenclature

Rule 10. Sect. 3. Publication of Names

In preparation for this Congress Vuillemin (1913) published a
paper in which he discussed bacterial classification and nomenclature.
He concluded that die best approach was to determine which generic
names were correct, to publish these names as geyiera conserifanda
and date all bacterial classification from 1915 when the list was to
have been adopted by the next Congress. A study of Vuillemin's
paper reveals that he himself does not formulate tenable bases for
differentiation of genera and his reasons for choosing certain generic
names and abandoning others will scarcely withstand critical analysis.

The Committee on Classification of the Society of American Bac-
teriologists suggested that the publication of the third edition of
Zopf's Die Spakpilze in 1885 be made the point of departure.

The first International Congress of Microbiology 1930 approved
the following statement:

In view of the adequate provisions made for special regulations

relating to the bacteria, and the feasibility of designating genera
conservanda among the bacteria by international agreement, it is
believed that the greatest stability will be conferred by the adoption

of the publication of Species Plantariim by Linnaeus in 1753 as the
point of departure for bacterial nomenclature. The adoption of this
date is recommended.

This recommendation was presented to the London Botanical
Congress in 1930 and approved. Through some oversight this date
was not included in the Botanical Codes published in 1935 and 1952.

There apparently are no names of organisms now included among
the bacteria that were proposed before 1773 (by Mueller) . Before
the year 1825 only two genera (Polyangium and Serratia) now in-
cluded in the bacteria were described as plants, and before 1850 only
three additional {Beggiatoa, Leptothrix and Sarcina) . All other
genera named were, until 1857, included in the Animal Kingdom.

The fixing of the date 1753 ensures that all published bacterial
names must be considered.

In zoology the corresponding beginning date of valid publication
is that of the tenth edition, 1758, of the Systema Naturae of Linnaeus.
This is fixed as the work that inaugurated the consistent application
of "binominal nomenclature" in zoology. This date of publication
is accepted as the starting point of zoological nomenclature and of the
Law of Priority. Any names of taxa in zoology published in other works
in 1758 are to be considered as published after the tenth edition of the
Systemae Naturae.

Bacteria and Viruses 61

Rule 11. SecN 3. Publication of Names

RULE 11. Publication is effected, under this Code, by sale or distribution of
printed matter to the general public or to bacteriological institutions. No
other kind of publication is accepted as effective (effective publication);
communication of new names at a public meeting or scientific conference
does not constitute effective publication.

Where reprints or separates from periodicals or other v^^orks are placed
on sale or issued in advance, the date on the separate is accepted as the
date of effective publication unless there is evidence that it is erroneous.

The date of acceptance of an article for publication as given in a publi-
cation does not indicate the effective date of publication and has no signifi-
cance in determination of priority of publication of names.

ANNOTATIONS

Rule 11. The text of the Botanical Code (Art. 29) is similar. The
reference to influence of date of acceptance on date of publication
Art. 30 states:

The date of effective publication is the date on Avhich the printed
matter became available as defined in Art. 29.
The Art. 29 reads:

Publication on and after 1 Jan. 1953 of a new name in tradesmen's
catalogues or in general and non-scientific newspapers, even if ac-
companied by a Latin diagnosis, does not constitute effective publi-
cation.
There is also a pertinent Recommendation:

Botanists and others are urged scrupulously to avoid publishing
new names or descriptions in ephemeral publications, in popular
periodicals, in any publication unlikely to reach the general botani-
cal public, in those produced by such methods that their penna-
nence is unlikely, or in abstracting journals.
Publication of names in botany cannot be effected by the issue of
microfilm made from manuscripts, typescripts or other unpublished
material.

Zoological Rules. The zoologists have distinguished between
publication before 1951 and after 1950. The designation "duly pub-
lished" is used apparently as the equivalent to "effectively published"
in the Bacteriological and Botanical Codes. A name is duly pub-
lished before 1951 only if it meets two requirements:

(1) The document containing the name must have been repro-
duced by printing or other mechanical method that ensured
that every copy is identical with every other copy.

(2) The document must have been issued for purposes of record
and consultation, and not for the sole consideration by special
persons, nor for particular purposes, nor for a limited time.

The above requirements must be met by a name published after
1950, but, in addition, the publication must have been reproduced
with ink on paper sufficiently durable to offer reasonable prospect
of permanence.

62 International Code of Nomenclature

Rule 11. Sect. 3. Publication of Names

The Zoological Rules recognize more clearly than either the
Botanical or Bacteriological Codes that there are numerous mechani-
cal methods of duplication other than by printing. It is, however,
recommended that publication, other than by printing, of material
affecting the status of a name should be avoided. In case of doubt,
the question is to be referred to the Commission for decision.

The Zoological Rules specify eight actions that do not constitute
publication. Some duplicate essentially the provisions of the other
Codes. Of special interest is that "Distribution of separates before
issuance of the book or paper to which they appertain" does not
constitute publication. Further, it is recommended that separates
"should not be distributed before the publication of the book or
serial to which they appertain."

Publication in zoology is not effected by distribution of such
duplicated material as microfilm or microcards.

It is recommended that whenever an author publishes a new
name for a nominal genus, subgenus, species or subspecies, he should
send notice promptly to a literature-recording serial (e.g.. Zoological
Record) .

The Zoological Rules contain numerous additional provisions
relating to publication. Many of these will be noted imder other
headings.

Bacteria and Viruses 63

Rule 12a. Sect. 3. Publication of Names

RULE 12a. A name of a taxonomic group (taxon) is not validly published
unless it is both (1) effectively published (see Rule 11) and (2) accompanied
by a description of the taxon or by a reference to a previously and effectively
published description of it.

NOTE. The words "valid" and "validly published" as used in this Code
mean "with standing in nomenclature", and the words "invalid" or "not
validly published" mean "without standing in nomenclature."

ANNOTATIONS

Rule 12a. The Botanical Code adds the parenthetical phrase " (direct
or indirect) " following the word "reference."

The Botanical Code also sitates that a combination (i.e., the name
of a species or of a taxon of subspecific rank) is not validly pub-
lished unless "the author definitely indicates that the epithet or epi-
thets concerned are to be used in that particular combination."

Botanists have added a rule which requires that, on or after
1 Jan. 1953, a new transfer or combination to be validly published
must have its basionym* (the name-bringing or epithet-bringing
synonym) clearly indicated with full reference to author and original
publication.

Botanists have a long tradition of requiring for valid publication
Latin diagnoses of newly described laxa of plants. At various times
unsuccessful attempts were made to change the rule. However, there
were many botanists who ignored the rule, particularly those who
described fungi, algae and bacteria. Final agreement was reached
that names of taxa published in languages other than Latin before
1 Jan. 1935 may be regarded as validly published. However, the
Botanical Congiess of 1930 approved the recommendation of the
Microbiological Congress of 1930 that new names of taxa of bacteria
are excepted from the general requirement. It was later agreed that
new names of algal taxa published before 1 Jan. 1958 do not require
Latin diagnoses (Botanical Code Art. 34, 1956) .

These rules are of interest to all microbiologists in that valid
publication of new names of bacterial and protozoan taxa may have
the diagnoses in any language, all new names of fungal taxa must
have diagnoses in Latin to be validly published, and after 1 Jan.
1958 the diagnoses accompanying new names of algal taxa must be
in Latin. Further, a new name of an algal taxon is validly published
after 1 Jan. 1958 only when accompanied by an illustration or figure
showing the distinctive morphological features.

* Also spelled basonym.

64 Infernafional Code of Nomenclature

Rules 12a, 12b. Sect. 3. Publication of Names

Throughout biolog\ the names given to all taxa must be Latin
or Latinized to be recognized as validly published.

The Botanical Code (Art. 35) also requires that publication on or
after 1 Jan. 1958 of the name of a new taxon of recent plants of the
rank of order or beloAv is valid only when the nomenclatural type
is indicated.

RULE 12b. Valid publication is not effected by placing a name on a culture
or preparation of bacteria in a collection or exhibit open to the public.

ANNOTATIONS

Rule 12b. New names of microorganisms have occasionally been
placed upon cultures, as in Type Culture Collections, and have some-
times been printed in catalogues, without descriptions. Such names
and new names placed upon museimi specimens for public exhibition
are not validly published.

In botany the placing of a new name upon a herbarium specimen
(exsiccatum) , even if accompanied by a description, does not consti-
tute valid publication.

Publication of a new name in zoology is not effected by placing
the name on the label of a museum specimen.

Bacteria and Viruses 65

Rule 12c. Sect. 3. Publication of Names

RULE 12c. A name (1) which is not accepted by the author who published
it, or (2) which is merely proposed in anticipation of the future acceptance
of the taxon concerned, or of a particular circumscription, position or rank
of the taxon, or (3) which is merely mentioned incidentally is not validly
published.

NOTE. Number (1) above does not apply to names or epithets published
with a question mark or other indication of taxonomic doubt, yet published
and accepted by the author. By "incidental mention" of a new name or
combination is meant mention by an author who does not intend to intro-
duce the new name or combination concerned.

ANNOTATIONS

Rule 12c (1). Beijerinck {Arch, neerl. d. sc. exactes, 1903, Sec. 2.
S:2\l) mentioned in a footnote to the publication in which he named
the genus Azotobacter that Parachromatium might be preferable to
show relationship to the genus Chromatium. It was not later used
by the author and may be regarded as not validly published.

De Petschenko {Zentr. f. Bakt. Abt. 1. Orig. 55:90-92. 1910) de-
scribed a parasitic infection of a protozoan by a spiral organism. In
a footnote he suggested the generic name Muellerina with the species
M. paramecii. Later {Arch. Protist. 22:248-298. 1911) he published
a much more detailed description, repudiated his previously sug-
gested names, and proposed the new names Drepanospira and D.
muelleri. These latter, rather than the former, have been accepted
by action of the Judicial Commission as validly published.

However, the author of a validly published name of a taxon may
not later change the name except to bring it into conformity with
the rules.

The Zoological Rules state that when a name is given it is pre-
sumed to be published for use in zoological nomenclature. However,
if the author of the name makes clear that the name is not intended
tor such use, it has no status either for priority or for homonymy.

66 International Code of Nomenclature

Rule 12c. Sect. 3. Publication of Names

Rule 12c (2) . Certain :iuthors have desired to publish a symmetrical
classification of bacteria, including generic names and descriptions
for jDurely hypothetical genera. For example, Fischer (Jahrb. wiss.
Bot. 27:4\. 1895) proposed the generic name Bactrinium to include
monotrichous, rod-shaped bacteria which produce endospores. The
author included no species, and none has been subsequently de-
scribed. The name was obviously proposed in anticipation of possible
future acceptance and it was not validly published.

The Botanical Code contains a similar provision, and states
further that if, on or after 1 Jan. 1953, two or more different names
are proposed simultaneously for the same taxon by the same author,
none of them is validly published.

A Zoological Commission has published the following pertinent
commentary on "The Nature of a Systematic Name":

...The Commission is unanimously of the opinion that a name,
in the sense of the Code, refers to the designation by which the
actual objects are known. In other words, we name the objects
themselves, not our conception of said objects. Names based upon
hypothetical forms have, therefore, no status in nomenclature and
are not in any way entitled to consideration under the Law of
Priority.

Bacteria and Viruses 67

Rules 12d, 12e, 1 2f. Sect. 3. Publication of Names

RULE 12d. A name of a taxon is not validly published when it is merely
cited as a synonym.

RULE 12e. The name of a taxon is not validly published by the mere mention
of the subordinate taxa included in it.

RULE 12f. The date of a name or of an epithet is that of its valid publica-
tion. For purposes of priority, hov/ever, only legitimate names and epithets
published in legitimate combinations are taken into consideration. In the
absence of proof to the contrary, the date given to the work containing the
name or epithet must be regarded as correct.

ANNOTATIONS

Rule 12d. Trevisan {Rendiconti Real. 1st. Lombard, d. Sci. e Lett.,
Ser. 2, 1879, 72:144) cited Malleomyces equestris Hallier as a synonym
of Micrococcus equestris which he regarded as the causal organism of
glanders. Inasmuch as all of Hallier's species were based upon mixed
cultures and his names, therefore, illegitimate (see Rule 24g) , this
incidental citation as a synonym by Trevisan does not validate the
name. Malleoyyiyces must date as a generic name from its proposal
by Pribram in 1933 {Klassification des Schizomyceten, p. 93) .
Rule 12f. Chondromyces crocatus Berkeley and Curtis 1857 (in
Berkeley, Introduction to Cryptoga^nic Botany, p. 313) is a name
appended to an illustration without description. The description
was published later (Berkely, Grevillea, 1874, 5:64) and valid publi-
cation was of the latter date.

68 International Code of Nomenclature

Recommendations 12a, 12b, 12c, 1 2d, 1 2e.

RECOMMENDATION 12a. When names of new taxa are published in works
written in a language unfamiliar to the majority of workers in bacteriology,
it is recommended that the authors publish simultaneously the diagnoses in
a more familiar language.

RECOMMENDATION 12b. Authors should indicate precisely the date of their
works. In the case of a work appearing in parts, the last published sheet
of the volume should indicate the precise dates on which the different fas-
cicles or parts of the volume were published as well as the number of pages
in each.

RECOMMENDATION 12c. When works are published in periodicals, the
author should require the publisher to indicate on the separates or reprints
the date (year and month, if possible the day) of publication and also the
title of the periodical from which the work is extracted. Separates or reprints
should always bear pagination of the periodical of which they form a part;
if desired, they may also bear a special pagination.

RECOMMENDATION 12d. An author who describes and names a new taxon
should indicate the rank of the taxon concerned and also the name and rank
of the next higher taxon, (i.e., the name of the family to which a new genus
is allocated, the name of the order in which a new family is placed).

RECOMMENDATION 12e. An author, when publishing the name of a new
taxon, should designate the type and, in the case of a species or subspecies
cultivable on artificial media, or otherwise preservable in pure culture in the
living state, indicate the collection where the type strain or its equivalent is
deposited.

ANNOTATIONS

Recomynendation 12e. In Botany a new name of a taxon of rank of
order or below published after 1 Jan. 1958 is valid only when the
nomenclatural type is designated. If the nomenclatural type is a speci-
men the place where it is conserved should be indicated.

The zoologists have a Rule (not a Recommendation) which re-
quires that a new generic or subgeneric name proposed subsequent to
31 Dec. 1930 be accompanied by a clear and unambiguous designation
of a type species for validation. This rule indirectly provides that no
new generic or subgeneric name is validly published if contained
species are not designated or indicated. The Zoological Code also
recommends that the author of the name of a new genus or subgenus
should not only indicate but expressly designate the type species by
name and, where the name of the species has previously been pub-
lished, he should cite the bibliographic reference and its new species
name (binomen) .

Bacteria and Viruses 69

Recommendation 1 2f. Publication of Names

RECOMMENDATION 12f. An author, when publishing a new generic or sub-
generic name, should give its etymology and also that of a new epithet when
the meaning is not obvious.

ANNOTATIONS

Recommendation 12f. The Botanical Code includes a similar Recom-
mendation. The Zoological Rules recommend that the author of a
generic name "based on a Greek or Latin ^vord" should indicate the
gender of the name and its etymology.

Compliance with this recommendation on the part of authors is
most helpful in many cases. The fact that "noveboracensis" means
"of New York" is not readily discovered by reference to a Latin dic-
tionary; one must know that the Latin name for York, England,
was Eboracum and that New York is Novum Eboracinu. Similarly,
knowledge that "Jersey" is derived from the older Latin name Caes-
area clears the meaning of "novaecaesareae" as "of New Jersey."

70 International Code of Nomenclature

Rule 13. Sec. 3. Publication of Names

RULE 13. The name of a genus or of a subgenus is not validly published
unless it is accompanied

(a) by a description of the genus or subgenus, or

(b) by citation of a previously and effectively published description of
the genus or subgenus, or

(c) by citation of a previously and effectively published description of
the genus as a subgenus or of the subgenus as a genus, or

(d) in the case of a monotypic new genus based on a new species, by a
combined generic and specific description, or

(e) by a description of one or more species or by citation of a previous
and effectively published description of one or more species included
in the genus.

NOTE. A description of a new species assigned to a monotypic new genus
is treated also as a generic description if the genus is not described.
Similarly a description of a monotypic new genus based on a new species
is treated also as a specific description if the generic name and specific
epithet are published together and the species is not separately described.

RECOMMENDATION 13a. A combined generic and specific description
should mention the points in which the new genus differs from related
genera.

ANNOTATIONS

Rule 13 (a) . The publication of the generic name Coryyiebacterium
Lehmann and Neumann (Bakt. Diag. 2:390. 1896) included a de-
scription of the genus and hence met one criterion for valid publication.
The publication of the subgeneric name Sarcinococcus Breed
(Bergey's Manual Det. Bad. Ed. 6, p. 285) included a description
of the subgenus and thus met one of the criteria for valid publication.

Rule 13 (b) . The publication of the generic name Rhodosphaera
Buchanan {Jour. Bact. 3:472. 1918) included a citation to a de-
scription of the genus Rhodococcus Molisch {Die Purpurbakterien
p. 20, 1907) , which generic name, though validly published, is an
illegitimate later homonym of Rhodococcus Zopf (1891) . The new
generic name Rhodosphaera proposed thus met a requirement for
valid publication.

Rule 13 (d). Beijerinck {Cent. Bakt. Abt. 2. 77:593. 1904) published
a combined generic and specific description based on the new species
Thiobacillus thioparus and thereby met the requirements of valid
publication of the generic name.

Bacteria and Viruses 71

Rules 14a, 14b. Sec. 3. Publication of Names

RULE 14a. The name of a species is validly published only when its publi-
cation conforms to the following requirements:

(1) It must be published as a binary combination consisting of a generic
name followed by a single specific epithet.

(2) It must be accompanied by a description of the species or by citation
of a previously and efFectively published description.

RULE 14b. The name of a subspecies (variety) is validly published only when its
publication conforms to the following requirements:

(1) It must be published as a ternary combination consisting of a generic
name followed by a single specific epithet and this in turn by a
single subspecific epithet.

(2) It must be accompanied by a description of the subspecies or by
citation of a previously and effectively published description.

(3) The author must clearly state or indicate that a subspecies is being
named.

ANNOTATIONS

Rule 14a. This rule in effect states that to be validly published and
hence to have standing in bacteriological nomenclature the name of
a species must conform to the requirement of Rule 6 that the name
of the genus shall be followed by a single specific epithet. If followed
by more than one specific epithet the species name is not a binary
combination, does not conform to the binomial system of nomen-
clature, and is not validly published. A specific epithet, as noted
under the Annotations of Rule 6, may consist of two or more related
words which should be written solid or hyphenated.

Rule 14b. This rule is intended to obviate the confusion resulting
from the naming of many microorganisms in the past. In some cases
it is very difficult, if not impossible, to determine whether an author
is proposing an illegitimate ternary combination as the name of a
species or a legitimate ternary combination as the name of a sub-
species. When a species is divided into subspecies, the subspecific
epithet of the name of the subspecies which includes the type of the
species must be the same as the specific epithet. The rule requires
that an author, when he names a new subspecies, make clear that the
ternary combination is that of a subspecies. (See Annotations under
Rule 7.) One of the difficult tasks of the systematic bacteriologist in
the future is to determine in the older literature of bacteriology the
legitimate names of subspecies, and to discard the illegitimate ternary
combinations used for names of species. Many cases will of necessity
be reviewed by the Judicial Commission and Opinions rendered in
order to remedy nomenclatural confusion.

Section 4

Citation of Authors and Names

RULE 15a. For the indication of the name (unitary, binary, or ternary) of a
taxon to be accurate and complete, and in order that the date may be
readily verified, it is necessary to cite the author who first published the
name in question. An alteration of the diagnostic characters or of the cir-
cumscription of a group without exclusion of the type does not warrant the
citation of an author other than the one who first validly published the
name.

RULE 15b. A name conserved so as to exclude its type is not to be ascribed to
the original author with such expressions as emend., mutatis choract., etc.; but
the author whose concept is conserved must be cited as authority.

NOTE. Retention of a name in a sense which excludes the type can be ef-
fected only by conservation.

NOTE. See Provisions 4 and 5 for method of conserving names.

ANNOTATIONS

Rule 15a. Examples: Plocamobacteriales Pribram (or Pribram 1933) ;
Proteus Hauser (or Hauser 1885) ; Serratia marcescens Bizio (or Bizio
1823).

Two or more authors may join in the publication of names of
new taxa. It is often awkward and unnecessary to cite the names of
all the co-authors, especially when there are more than two. It has
become increasingly common to cite the name of the first author fol-
lowed by et al. For example, Hauduroy, Ehringer, Guillot, Magrou,
Prevot, Ronet and Urbain, in their Dictionnaire des Bacteries Patho-
genes, have proposed a large number of new combinations (comb,
nov.) . To list seven names after each of these new combinations is
obviously undesirable. The formula Pasteurella ellingeri Hauduroy
et al. has been used.

In the first and second editions of Bergey's Manual of Determina-
tive Bacteriology, the five members of the Committee on the Manual
(Bergey, Harrison, Breed, Hammer and Huntoon) have been re-
garded as joint authors of all new names of taxa included. Citation
is customarily to Bergey et al.

[73]

74 International Code of Nomenclature

Rules 15a, 15b. Citation of Authors and Names

One of the most common errors in bacterial nomenclature is the
ascription of scientific names to individuals who did not propose
them. For example, Lauda (1930) in his section on Bartonella in
the Kolle, Kraus and Uhlenhuth "Handbuch der pathogenen Mikro-
organismen" uses the ascription "Bartonella bacilliformis Barton
1909." What Lauda intended to indicate, was that Barton discovered
or described the organism. The correct ascription in this instance is
Bartonella bacilliforTuis Strong, Tyzzer, Brues, Sellards and Gastiaburu,
1915 or, more briefly, Bartonella bacilliformis Strong et al., 1915. This
error of citing the discoverer or describer of an organism, rather than
the author of its name, has led to much unnecessary confusion .
The Zoological Code includes a helpful definition of "author."
The author of a scientific name is that person who first published
the name in connection with an indication, a definition, or a de-
scription, unless it is clear from the contents of the publication that
some other person is responsible for said name and its indication,
definition, or description.

Bacteria and Viruses 75

Recommendations 15a, 15b. Citation of Authors and Names

RECOMMENDATION 15a. When the alteration of diagnostic characters or
circumscription of a taxon has been considerable, the nature of the change
and the author responsible may be indicated by adding suitable abbreviated
words such as emend, (emendavit), mut. char., excl. gen., excl. spec, etc.

RECOMMENDATION 15b. When a name with a description or reference to
a description by one author is published in a work of another author, the
word in should be used to connect the names of the two authors.

ANNOTATIONS

Recommendation 15a. Example: Bacillus Cohn 1872 emend. Migula
1894.

Recommendation 15b. Example: Simonsiella mueUeri Schmid in
Simons 1922.

76 International Code of Nomenclature

Rule 16. Sect. 4. Citation of Authors and Names

RULE 16. When a genus, a subgenus, a species, or a subspecies (variety) is
altered in rank but retains its name or epithet, the original author must be
cited in parentheses, followed by the name of the author who efFected the
alteration. The same holds when a subgenus, a species, or a subspecies
(variety) is transferred to another genus or species with or without altera-
tion of rank.

ANNOTATIONS

Rule 16. Example: Philip 1943, proposed the name Coxiella for a
new subgenus of the genus Rickettsia. The name of the subgenus
with the author citation is Coxiella Philip, or Coxiella Philip 1943.
Philip elevated the subgenus to the rank of genus, the generic name
with author citation is Coxiella (Philip) Philip 1948. An organism
regarded as the cause of syphilis was named Spirochaete pallida; with
author's citation it is Spirochaete pallida Schaudinn and Hoffmann
1905. Vuillemin concluded that the organism was improperly placed
in the genus Spirochaete and proposed the generic name Spironeyna
with the species name Spironeina pallidum, with author citation,
Spironeyna pallidum (Schaudinn and Hoffmann 1905) Vuillemin 1905.
Schaudinn noted that Spironeyna was illegitimate because it is a later
homonym and proposed the new generic name TrepoJiema; the
species name with author citation is Treponema pallidum (Schaudinn
and Hoffmann 1905) Schaudinn 1905.

Botanical Code. The Rule is essentially the same, but is more
explicit in requiring that the name or epithet published by the
original author was legitimate.

Zoological Rules. The rules are somewhat more explicit. When
it is desired to cite the name of the author of the name of a taxon.
the citation should follow the scientific name without separation by any
mark of punctuation.

When a species or subspecies is transferred to another than the
original genus or when the specific name (specific epithet) is combined
with any other generic name than that with which it was originally
published, the name of the author of the specific name (specific
epithet) is retained in parentheses. Citation of the author of the new
combination is desirable; his name follows the parentheses. In the
Zoological Code rather more emphasis is laid ujx)n the citation of the
name of the author of the specific epithet, in the Botanical and Bac-
teriological Codes rather more upon the author of the combination.

Abbreviations of the names of authors. There is no provision in
the Bacteriological Code for the abbreviation of the names of authors
of scientific names. In general such abbreviations are unnecessary
and inadvisable.

Bacteria and Viruses 77

Rule 16. Citation of Authors and Names

The Zoological Code recommends that in citation ot the name
of an author there should be no abbreviation unless the author be
deceased and of such importance that the abbreviation would be
readily recognized.

The Botanical Code differs from the Bacteriological and Zo-
ological Codes in that it emphasizes the use of abbreviations of the
names of authors. It specifically recommends such abbreviation and
suggests standard methods of abbreviating names. This custom of
abbreviation has come about in part because certain individuals gave
a large number of names. That the abbreviation L. means Linnaeus
and DC, A. P. De Candolle, is generally recognized.

78 International Code of Nomenclature

Recommendations 16a, 16b, 16c, 16d, 16e. Citation of Authors and Names

RECOMMENDATION 16a. When citing a name published as a synonym, the
words "as synonym" or "pro synon." should be added to the citation.

When an author has published as a synonym a name from a manuscript
of another author, the word ex should be used to connect the names of the
two authors.

RECOMMENDATION 16b. When citing in synonymy a name invalidated by
an earlier homonym, the citation should be followed by the name of the
author of the earlier homonym preceded by the word "non", preferably with
the date of publication added. In some instances, it will be advisable to cite
also any later homonym or homonyms.

RECOMMENDATION 16c. In the citation of a nomen nudum, its status should
be indicated by adding nom. nud.

RECOMMENDATION 16d. If a generic name antedated by one of its synonyms
or by a homonym is accepted as a nomen conservondum, the abbreviation
nom. cons, should be added to the citation.

RECOMMENDATION 16e. Names cited in synonymy should be spelled
exactly as published by their authors. If any explanatory words are re-
quired, these should be inserted in brackets. If a name is adopted with
alterations from the form as originally published, it is desirable that in full
citations the exact original form should be appended.

ANNOTATIONS

Recommendation 16a. This is essentially the same as the correspond-
ing Recommendation in the Botanical Code. Search has thus far
revealed no good example in bacteriology. That given in the Botani-
cal Code may illustrate.

Example: Myrtus serratus is a species name used in a manuscript
by Koenig. It was published by Steudel as a synonym of Eugenia
laurina Willd. It should be cited as Myrtus serratus Koenig ex
Steudel, Nomencl. 321 (1 82 1) pro. syn.

Recommendation 16b. Thaxter (1892) published the name Myxo-
coccus for a genus of bacteria. Gonnerman (1907) later proposed
Myxococcus as the name of another genus of bacteria. The name
given by Gonnerman should be cited as: Myxococcus Gonnerman
1907 non Thaxter 1892.

Guillebeau 1890 proposed the generic name Chlorobacterium;
Lauterborn 1915 proposed the same name for another genus of bac-
teria. The citation of the latter should be Chlorobacterium Lauter-
born 1915 non Chlorobacterium Guillebeau 1890.

Bacteria and Viruses 79

Recommendations 1 6c, 1 6d, 1 6e. Citation of Authors and Names

Recommendation 16c. A nomen nudum (naked name) has been de-
fined in zoology as a name published without any definition, de-
scription or indication of the taxon to which it is intended to apply,
and hence without status in nomenclature.

Example: Fischer 1895 proposed the generic name Clostrinium to
include rod-shaped bacteria with a single polar flagellum and with
cells which become spindle-shaped when sporulation occurs. No
species was named by the author and none has since been described.
It may be regarded as a nomen nudum.

Recommendation 16d.

Example: Guillebeau 1890 proposed the name Chlorobacterium for
a bacterial genus. The generic name Pseudomonas Migula 1894, a
later synonym of Chlorobacterium Guillebeau, was designated as
a nomen generis conservandum (conserved name of a genus) , and
the generic name Chlorobacterium Guillebeau placed in the list of
nomina generum rejicienda (rejected names of genera) . A correct
citation would be Pseudomonas Migula 1894 nom. cons.

Recommeyidation 16e. Authors sometimes spell names of taxa dif-
ferently from the original or the accepted spelling. In some cases the
change is intentional. For example, Enderlein 1917 preferred the
spelling Corynobacterium of the generic name Corynebacterium
Lehmann and Neumann 1896. In a list of synonyms each name should
be spelled exactly as published by its author. Freqtiently the word sic
in parentheses is appended to the citation to indicate that the spell-
ing given is that of the author and not a misprint.

Changes in Names
as a Result of Segregation, Union, or Change in Rank of Taxa

RULE 17a. An alteration of the diagnostic characters or of the circum-
scription of a taxonomic group (taxon) does not warrant a change in its
name, except as this may be necessitated by transference of the taxon, by
its union with another taxon of the same rank (Rule 19), or by a change of
its rank (Rule 22).

RULE 17b. When a genus is divided into two or more genera, the generic name
must be retained for one of them, or (if it has not been retained) must be
re-established. When a particular species was originally designated as the
type, the generic name must be retained for the genus including that species.
When no type was designated, a type must be chosen.

The same rule is applied when a subgenus is divided into two or more
subgenera.

RULE 17c. The name of a taxonomic group must be changed if the nomen-
clatural type of the taxon is excluded.

ANNOTATIONS

Rule 17a. There is no warrant for changing the name of a taxon
because the boundaries (circumscription) of the taxon are changed.
The addition of new species to a genus may require a modification
of the description (tabulation of diagnostic characters) of the genus,
but not a change of name. For example, the genus Pseudomonas
Migula 1894 is defined by some authors to include polar flagellate
plant pathogens with yellow, non-diffusing pigment; others use the
generic name Xanthomonas Dowson 1939 for the latter. Contraction
or expansion of the boundaries of the genus itself is no reason for
changing the name as long as the type species is retained within the
genus.

Rule 17b. Example: When Donker (1926) proposed and named a
new genus Aerobacillus to include those species of the genus Bacillus
which are aerogenic, he correctly retained the generic name Bacillus
for that portion which included the type species Bacillus subtilis
Cohn. Winslow and Rogers (1906) divided the genus Staphylococcus
Rosenbach (1884) into two genera named Aurococcus and Albo-
coccus. One of the genera should have retained the name Staphylo-
coccus, that containing the type species.

The corresponding Article in the Botanical Code has essentially
the same wording.

The Zoological Code provides that when two or more restricted
genera are formed by the division of a genus, the available name

[81]

82 International Code of Nomenclature

Rules 17b, 17c. Changes in Names of Taxa

must be retained for one of the genera, that containing the type
species if one had been originally established.

Rule 17c. The Enterobacteriaceae Subcommittee in its report pre-
sented to the Rome Congress (1953, Int. Bulletin, •/:47) included
Aerobacter aerogenes in the genus Klebsiella; this action deprived
Aerobacter Beijerinck of its generally accepted type species. Edwards
and Fife (1955, /. Bact., 70:^82) proposed that Aerobacter should be
redefined with Aerobacter cloacae as type species. To become effective
this would require international approval. Later, Hormaeche and
Edwards (Internatl. Bull. Bact. Nomen. and Tax. 5:111-115. 1958)
withdrew the earlier proposal and substituted for it a redefinition of
the genus Aerobacter based on a motile strain which they designated
A. aerogenes.

Bacteria and Viruses 83

Rules 18a, 18b. Sect. 5. Changes in Names of Taxa

RULE 18a. When a species is divided into two or more species, the specific
epithet must be retained for one of them, or (if it has not been retained)
must be reinstated. When a type has been designated for the species, the
specific epithet must be retained for the species including that type. When
no type was designated, one must be chosen.

The same rule applies to subspecies (varieties); for example, to a sub-
species (variety) divided into two or more subspecies (varieties).
RULE 18b. When a species is transferred to another genus (or placed under
another generic name for the same genus), without change of rank, the
specific epithet must be retained or (if it has not been retained) must be re-
established unless one of the following obstacles exists: (1) the resulting
binary name is a later homonym or tautonym or (2) there is available an
earlier validly published specific epithet.

When the specific epithet, on transference to another generic name, has
been applied erroneously in its new position to a difFerent species, the new
combination must be retained for the organism on which the epithet was
originally based.

ANNOTATIONS

Rule 18a. Example: Frank (1890) named Rhizobhim leguminosarum
as the single species of the genus to include the bacteria responsible
for nodulation of the roots of leguminous plants. Baldwin and Fred
(1929) recognized several species described from the roots of different
legumes. They rightly retained the name Rhizohium leguminosarum
Frank as the type species to include the bacteria from species of
Lathyrus and related genera.

The rule in botany is essentially the same.

The corresponding Article in the Zoological Rules states that
when a nominal species (or a subspecies) is divided into two or more
restricted species (or subspecies) the division is governed by the
same Rules as in the division of a genus.

84 International Code of Nomenclature

Rule 19, Recommendation 19a. Changes in Names of Taxa

RULE 19. When two or more taxa of the same rank are united, the oldest
legitimate name or (in species and subspecies) the oldest legitimate epithet
is retained. If the names or epithets are of the same date, the author who
first unites the taxa has the right to choose one of them, and his choice
must be followed.

RECOMMENDATION 19a. Authors who have to choose between two generic
names should note the following recommendations: (1) Of two names of
the same date, to prefer the one which was first accompanied by the de-
scription of a species. (2) Of two names of the same date, both accompanied
by descriptions of species, to prefer the one which, when the author makes
his choice, includes the larger number of species. (3) In cases of equality
from these various points of view, to prefer the more appropriate name.

ANNOTATIONS

Rule 19. The genera Micrococcus Cohn 1872 and Staphylococcus
Rosenbach 1884 were united by Hucker who correctly chose the
generic name Micrococcus as having priority.

The corresponding Article of the Botanical Code is essentially
the same.

The Zoological Rules provide that when two or more genera or
subgenera are united, the genus formed must take the oldest available
generic or subgeneric name of its components. The rule just quoted
applies as well when two or more species or subspecies are united to
form a single species or subspecies.

Recomviendation 19a. This Recommendation corresponds to that in
botany.

Bacferia and Viruses 85

Rules 20, 21. Sect. 5. Changes in Names of Taxa

RULE 20. When several genera are united as subgenera under one generic
name, the subgenus including the type of the generic name used must bear
that name unaltered.

ANNOTATIONS

Rule 20. Pederson 1945 included the genera Thermobacterium Orla-
Jensen 1919, Streptobacterium. Orla-Jensen 1919, and Betabacterium
Orla-Jensen 1919, in the genus Lactobacillus Beijerinck 1901. The
type species of Lactobacillus is L. caucasicus (Beijerinck 1889) Bei-
jerinck 1901. Pederson defined the subgenus Thermobacterium to
include L. caucasicus {Lactobacillus [Thermobacterium] caucasicus) .
This recognition of Thermobacterium as a subgenus does not con-
form to Rule 20. The subgenus which includes the type species of
the genus should bear the name Lactobacillus, of which the sub-
generic name Therjuobacterium (Orla-Jensen) Pederson becomes a
svnonvm.

RULE 21. When several species are united as subspecies (varieties) under
one specific name, the subspecies v/hich includes the type of the species used
must be designated by the same epithet as that of the species.

ANNOTATIONS

Rule 2L Smith 1948, in his revision of the species Bacillus subtilis
Cohn 1872, included as subspecies Bacillus aterrimus Lehmann and
Neumann 1896 and Bacillus niger Migula 1900. He correctly re-
named them Bacillus subtilis var. aterriynus and B. subtilis var. niger.
Under the Bacteriological Rule Smith's revision automatically cre-
ated a third variety (subspecies) Bacillus subtilis var. subtilis of
which the type cultuie is that of the species.

86 International Code of Nomenclature

Rules 22a, 22b, 22c. Sect. 5. Changes in Names of Taxa

RULE 22a. When a toxon of a rank higher than a genus and not higher than
an order is changed in rank, the stem of the name must be retained and
only the suffix altered (-inae, -eoe, -oideae, -oceoe, -ineae, -ales), unless the
name must be rejected under Section 6.

RULE 22b. When a subgenus becomes a genus, or a genus becomes a sub-
genus, the original name must be retained unless it is rejected under Section
6.

RULE 22c. When a subspecies (variety) becomes a species, or a species be-
comes a subspecies (variety), the original epithet should be retained unless
the resulting combination is rejected under Section 6.

ANNOTATIONS

Rule 22a. When a tribe is raised to the rank of a family, the suffix
-eae of the name must be changed to -aceae; for example, Pasteurelleae
becomes Pasteurellaceae. The new name must not contravene the pro-
visions of Rule 24.

Essentially the same provision is to be found in the Botanical
Code.

The Zoological Rules prescribe definitely only the suffixes to be
used for family and subfamily, namely, -idae and -inae. The basic
concept is quite different from that of bacteriology and botany. A
Family-Group of categories is recognized including all ranks between
genus and suborder. All names of the Family-Group of categories
are coordinate despite the endings of the names. For determination
of priority each name dates from its original publication for any
category within the family group. This priority is retained even
though treated as belonging to a higher or lower category within the
Family-Group. All Family-Group names are regarded as homonyms
if they are identical apart from their terminations.

Bacteria and Viruses 87

Rules 22b, 22c. Changes in Names of Taxa

Rule 22b. Under the Bacteriological Code the name of a genus is
not changed when it becomes a subgenus, nor is the name of a sub-
genus changed when it becomes a genus. This rule definitely fixes the
coordinate nomenclatural status of generic and subgeneric names.

The Botanical Code does not recognize the equal priority status
of generic and subgeneric names. The pertinent article (Art. 70)
states that when the rank of a genus or an infrageneric (all ranks
below genus) rank is changed, the correct name is the earliest legiti-
mate name available in the new rank; in other words, a name of a
genus or subgenus has no priority outside its own rank. An author
who raises a subgenus to the rank of genus need not adopt the sub-
generic name for the name of the new genus.

The Zoological Rule states that generic and subgeneric names are
subject to the same Rules and Regulations and are nomenclaturally
coordinate, thus resembling the Rule in the Bacteriological Code.
When a later author changes the rank of a genus to that of a sub-
genus, the name of the genus becomes the name of the subgenus.
Similarly a subgenus when raised to the rank of a genus retains its
name.

Rule 22c. Names of species and subspecies are subject to the same
Rules and Recommendations and are nomenclaturally coordinate.
When a species becomes a subspecies or a subspecies becomes a species
the epithet (specific or subspecific) is unchanged.

The Zoological Rules are essentially the same.

The Botanical Code on the contrary does not regard specific and
subspecific epithets as nomenclaturally coordinate and states that in
no case does an epithet have priority outside its own rank. An author
who raises the rank of a subspecies to that of a species is not com-
pelled to retain the subspecific epithet for that of the species.

Rejection and Replacement of Names

RULE 23. A legitimate name or epithet must not be rejected merely because
it is inappropriate, or disagreeable, or because another is preferable or
better known, or because it has lost its original meaning.

ANNOTATIONS
Rule 23. Neglect or misunderstanding of this rule has led in bac-
teriology to much nomenclatural confusion. It has been assumed by
some bacteriologists that all names given to taxa should be appropri-
ate or agreeable. But there is no rule sanctioning this interpretation
in botany, bacteriology or zoology. This is in line with the principle
that all three Codes aim at fixity of names. A name of any taxon is
basically an arbitrary symbol. The Codes do, however, offer guidance
on how best to coin appropriate names for new taxa. Bacteriological
Rules governing the formation of names of taxa higher than genus
require that the names be Latin or Latinized and in the plural num-
ber. Little use is made in bacteriology of names of taxa higher than
order. It is recommended (Recommendation 2a) that the names of
new orders and suborders be based upon the name of the type genus
of the type family. The names of other taxa between suborder and
genus must be formed (Rule 3) by the addition of the appropriate
suffix to the stem of the correct name of the type genus. The name
of a higher taxon based upon the name of its type genus cannot be
inappropriate. Nevertheless, there have been many objections raised
to the use of certain properly constructed family and other names
because they are allegedly inappropriate. These objections are based
upon fallacious reasoning. For example, an author concludes that the
genera Lactobacillus and Streptococcus are so closely related (have
so many characters in common) that they should be placed together
in a family. Under the rules the family name must be based upon
the name of the type genus. If the author chooses Lactobacillus as
the type genus, the family name under the rules becomes Lactobacil-
laceae, including the two genera Streptococcus and Lactobacillus. The
objection to this family name has been raised that the name Lacto-
bacillaceae means "resembling milk rodlets" and is inappropriate be-
cause the other genus. Streptococcus, of the family does not include
ix>ds. The fallacy of this reasoning is evident if consideration is given
to the true meaning of the family name Lactobacillaceae (Lacto-
bacillus-\ike) . It does not mean or imply that organisms belonging
to the family Lactobacillaceae must be rod-shaped or related to milk.
The genus Streptococcus has enough characters in common with the
genus Lactobacillus so that it may well be included in the same
family. The morphology is different, but in certain other significant
characteristics Streptococcus resembles Lactobacillus.

[89]

90 International Code of Nomenclature

Rule 23. Rejection and Replacement of Names

The name of a genus may be "inappropriate" when it is de-
scriptive of some character of one or more species of the genus, but
not descriptive of all contained species. A generic name implying
the presence of some color may be "inappropriate" when applied to
certain species of the genus. This "inappropriateness" is no adequate
reason for replacing the generic name by another.

In Recommendation 6b of the Bacteriological Code emphasis is
laid upon the desirability of naming a species by use of a specific
epithet "which, in general, gives some indication of the appearance,
the characters, the origin, the history, or the properties of the species."
A species name including such a specific epithet is appropriate. How-
ever, a Recommendation is a guide to good usage, but a name which is
not in accordance with the provisions of a Recommendation is not il-
legitimate and cannot be replaced on the basis of inappiopriateness.

The corresponding Article of the Botanical Code is essentially
similar.

The Zoological Code states that when once published, a name is
not to be rejected because of inappropriateness. A generic name such
as Apus ("that which is footless") as applied to birds with feet is
not to be rejected. One provision in zoology has no counterpart in
bacteriology or botany: no name shall have any status in nomen-
clature that is "reasonably" calculated to offend on "political, re-
ligious or personal grounds." Such a name may be suppressed if
upon application of any person to the International Commission it
is found offensive in any language.

Bacteria and Viruses 91

Rule 24. Sect. 6. Rejection and Replacennent of Names

RULE 24. A name must be rejected if it is illegitimate, i.e., if it is contrary
tc a Rule. The publication of an epithet in an illegitimate combination must
not be taken into consideration for purpose of priority.
A name of a taxon is illegitimate:

a. If it was nomenclaturally superfluous when published, i.e., if the
taxon to which it was applied, as circumscribed by its author, in-
cluded the type of a name which the author ought to have adopted
under one or more of the Rules.

b. If it is a binary or ternary name published in contravention of Prin-
ciple 9 and Rules 17-23, i.e., if its author did not adopt the earliest
legitimate epithet available for the taxon with its particular circum-
scription, position, and rank.

c. If its specific epithet must be rejected under Rule 25.

d. If it is a later homonym of the name of a taxon of bacteria, plants,
ov protozoa; that is, if it duplicates a name previously and validly
published for a taxon of the same rank based on a different type.
Even if the earlier homonym is illegitimate, or is generally treated
as a synonym on taxonomic grounds, the later homonym must be
rejected. When an author simultaneously publishes the same new
name for more than one group, the first author who adopts one of
them, or substitutes another name for one of them, must be followed.
NOTE: Mere orthographic variants of the same name are treated
as homonyms when they are based on different types.

e. If it is used with different meanings and so has become a long
persistent source of error. A list of names (nomina ambigua) to be
abandoned for this reason will be included under nomina rejiciendo.

f. !f its application is uncertain (nomen dubium). A list of names to
be abandoned for this reason will be included under nomina re-
jiciendo.

g. If the characterization of the group was based upon an impure or
mixed culture. A list of names to be abandoned for this reason
(nomina confuso) will be included under nomina rejiciendo.

h. If it was based upon an abnormality.

ANNOTATIONS

Rule 24(1. The publication of the generic name Dicrobactriim Ender-
lein 1917 was superfluous because of the publication of Serratia
Bizio 1823, based upon the same type species. Calym?natobacterinm
granuloinatis Aragao and Vianna 1912 has priority over Donovania
granulomatis Anderson, De Monbreun and Goodpasture 1945. The
latter name is superfluous.

Rule 24b. The specific epithet used in the name of a species must
be the earliest available under the rules. The specific epithet mar-
cescens of the species Serratia rnarcescejis Bizio 1823 was the first
given, and renders illegitimate the use of the epithet of Zaogalactina
imetroja Sette 1824, of the epithet in Protococcus imetrophus Mene-
ghini 1838, and of that in Monas prodigiosa Ehrenberg 1849.

92 International Code of Nomenclature

Rule 24. Rejection and Replacement of Names

Rule 24d. Pfeifferelln Buchanan 1918, proposed as a generic name
in bacteriology, is illegitimate because it is a later homonym of
Pjeifferella Labbe 1899, proposed as a name for a genus of protozoa.

Rule 24e. Bncterium Ehrenberg 1828 has as its type species Bacterium
triloculare Ehrenberg 1828. The type species has never been identified
or observed with certainty since Ehrenberg's first observation. The
generic name has been applied by later writers in so many ways as
to lead to major confusion and misunderstanding. The Judicial
Commission issued an Opinion placing the generic name Bacterium
Ehrenberg 1828 in the list of rejected names {nomina rejicienda) ,
as an ambiguous name (nomen ambiguum) .

Rule 24 j. Names have been given to many taxa of bacteria, particu-
larly to species, for which there are no descriptions adequate for
identification, and of which there are no known cultures. Such names
are placed in the list of noynina rejicienda as nojnina ditbia. Oc-
casionally a later student may be able to reisolate and identify ade-
quately an organism named in the list of nomiua duhia, from which
list it will then be removed.

Rtile 24g. There are numerous instances in the literature of bac-
teriology in which the description of a purported new species was
based upon a mixed or impure culture. Weinberg and Seguin (1918)
described a new species under the name of Bacillus aerofoetidus. This
was later shown to be a mixed culture containing three species, two
of them being Clostridium welchii (C. perjringens) and C. oede-
matis. The characters of the genus Malleomyces Hallier 1870 were
derived from various fungi and bacteria erroneously supposed to be
growth forms of a single organism. The name is illegitimate.

Rule 24h. A bacterial culture which became infected with a bac-
teriophage might produce colonies not characteristic of the species,
and the supposedly different organism might be given a new name.
Such a variant culture would be classed as an abnormality, and a
new species name gi\en to distinguish it would be illegitimate.

Bacteria and Viruses 93

Recommendation 24a. Sect. 6. Rejection and Replacement of Names

RECOMMENDATION 24a. Authors should avoid introducing into bacteriology
as generic names such names as are in use in zoology.

ANNOTATIONS

Recoynmendaiton 24a. The author of a new generic name in bacteri-
ology should make sure that a name he wishes to propose is not a
later homonym of another generic name in bacteriology, in botany
or in protozoology. He is concerned only with names that have been
validly published; others have no nomenclatural status.

In bacteriology there exists no adequate list of validly published
generic and subgeneric names. Such a list is in preparation and
should be published as an appendix to future editions of the Bac-
teriological Code. The International Bureau for Plant Taxonomy
and Nomenclature is preparing a card catalogue of validly published
names of plant genera, including the bacteria. As this list is pre-
pared the bacterial names will be published in the International
Bulletin of Bacteriological Nomenclature and Taxonomy. Until this
list is available, most questions of availability can be answered by
reference to the indices of the Seventh Edition of Bergey's Manual
of Determinative Bacteriology and its companion volume, the Index
Bergeyana. In obscure or difficult cases and determinations, authors
may request assistance from the Judicial Commission and its Edi-
torial Board.

A list of validly published names of plant genera is in prepara-
tion as noted above in the form of a card catalogue. Until this is
completed one should consult for generic names of flowering plants
and ferns the Index Kewensis with its numerous supplements. Ains-
worth and Bisby's Dictionary of the Fungi, Ed. 4, 1954, gives a
virtually complete list of the generic names of fungi pioposed up to
about 1953. The Index of Fungi (issued by the Commonwealth
Mycological Institute, Kew) lists generic names published since.

Becker prepared a list of generic names of protozoa taken from
the list of generic names of the Animal Kingdom (Schulze and
Kiickenthal, Neave) . The Zoological Record, with its carefully pre-
pared lists of names of taxa proposed in zoology, is probably the best
reference source in this field.

Botanical Code. The Rules of the Bacteriological Code relative
to rejection of names and epithets do not differ materially from those
of the Botanical Code on which they were originally based.

94 International Code of Nomenclature

Recommendation 24a. Rejection and Replacement of Names

One botanical rule not duplicated in the Bacteriological Code
relates to acceptance in botany of names of taxa transferred from the
animal to the plant kingdom. A name of a taxon is illegitimate if
it is the name of a taxon which at the time of such transfer becomes
a homonym of the name of an existing jjlant taxon. Names of taxa
transferred from the plant to the animal kingdom retain their botani-
cal nomenclatural status for purposes of homonymy.

That there will be transfer of taxa to or from branches of zoology
other than protozoology to botany is improbable.

Zoological Rules. A "Code of Ethics" which appeared in some
earlier editions of the Rules has been rephrased. A zoologist who
notes that a generic or specific name published by a living author as
new is a later homonym and therefore unavailable should notify the
author of the facts and give ample opportunity for a substitute name
to be proposed by him.

When an organism is transferred from the animal to the vege-
table kingdom, its generic name for purposes of homonymy retains
its status under the Zoological Code.

Bacteria and Viruses 95

Rule 25. Sect. 6. Rejection and Replacement of Names

RULE 25. A specific or subspecific epithet is illegitimate in the following
special cases and must be rejected:

a. When it is merely a word not intended as a specific epithet.

b. When it is merely an ordinal adjective used for enumeration.

c. When it exactly repeats the generic name (tautonym).

d. When it is a later homonym within the same genus, that is, when it
duplicates a specific epithet previously and validly published for a
species of the same genus based upon a different type.

e. When it was published in a work in which the Linnaean system of binary
nomenclature for species was not consistently employed.

ANNOTATIONS

Rule 23. Note that this rule governs only certain special cases. A
specific epithet is illegitimate also if it contravenes any rule of the
Bacteriological Code, that is, it is illegitimate if its author did not
use the earliest legitimate epithet available for the taxon being named.
Rule 23a. Authors sometimes do not completely identify by name
an organism described, but use an expression such as Clostridium sp.

(species) . The expresssion means that the author regards the or-
ganism as belonging to the genus Clostridium, but with species in-
determinate. The word species is "merely a word not intended as a
specific epithet." In some fields of biology, particularly in botany,
the pronominal adjective "qualis" (some kind of) , also "novus"

(new) , have been used in place of a specific epithet before a suitable
epithet has been decided upon. The use of qualis and novus in this
sense does not preclude their use if desired as specific epithets, as in
Bacterium qualis Steinhaus 1941 and Plectridium novum Huss 1907.

Rule 23b. Authors in some cases have used ordinal adjectives instead
of specific epithets, particularly in enumeration of several new species.
The sixth species of the genus Bacillus might be termed Bacillus
sextus. Sextus as here used is not a legitimate specific epithet. This
illegitimacy does not extend to the intentional employment of ordi-
nal adjectives as specific epithets. There is a Bacillus tertius Henry
1916 and a Clostridium sex turn Prevot 1940.

Letters of the Latin or Greek alphabet have sometimes been used
in the enumeration of new species and subspecies, as Bacillus a,
Bacillus h, etc. Letters thus used do not constitute legitimate specific
epithets, although specific epithets such as delta, alpha, etc. may be
legitimate. An "Opinion" relating to the Zoological Code states that
"serial letters, as a, b, c, etc. are not acceptable as specific names."

96 Infernational Code of Nomenclature

Rule 25. Rejection and Replacement of Names

Rule 25c. In bacteriology and botany it is the rule that a specific
epithet is illegitimate it it exactly repeats the name of the genus (i.e.,
if it is a tautonym) . When Castellani and Chalmers (1919) decided
to make Bacterium alcaligenes Lehmann and Neumann 1899 the
type species of a new genus Alcaligenes they were debarred from the
use of Alcaligenes alcaligenes and chose A. faecalis. The rule in
zoology approves, indeed encourages, the use of specific epithets
which are tautonyms.

Bacteria and Viruses 97

Rule 26. Sect. 6. Rejection and Replacement of Names

RULE 26. The name or epithet to be rejected according to Rules 23-25 is
replaced by the oldest legitimate name, or (in a combination) by the oldest
legitimate epithet which will be, in the new position, in accordance with the
Rules. If none exists, a new name or epithet must be chosen.

NOTE. Where a new epithet is required, an author may, if he wishes, adopt
an epithet previously given to the taxon in an illegitimate combination, if
there is no obstacle to its employment in the new position or sense; the
resultant combination is treated as a new name.

ANNOTATIONS

Rule 26. The Botanical Code has essentially the same wording.

This is the application of the Law of Priority, clearly stated in
Principle 9. Examples: The generic name Polyayigiiim Link 1809
replaced the generic name Myxobacter Thaxter 1892 when the earlier
name with description was found by Thaxter. Similarly the species
name Myxobacter aureus Thaxter 1893 became a synonym of Poly-
angiwn vitellinum Link 1809.

The species Bacterium aeruginosiun Schroeter 1872 was later in-
dependently named Bacillus pyocyaneus by Gessard 1882. Migula
proposed the new generic name Pseudomonas and recognized the
specific epithet having priority, in Pseudomonas aeruginosa (Schroe-
ter) Migula.

The Zoological Code under Law of Priority states that the name
to be used for a genus or for a species must be the oldest name that
fulfills three requirements. (1) The name must have been published
and have been accompanied by an indication, or a definition, or a
description of the taxon to which it appertains. (2) The author of
the name must have consistently applied the principles of binominal
nomenclature in the publication. (3) Any name published after
1930 must have had included in its publication a statement by the
author in which he attempted to indicate the distinguishing charac-
ters of the taxon. Further, if published as a substitute for a rejected
homonym, the publication must give a reference to the name re-
placed. If the name published is that of a genus or subgenus, it must
have been published with a type species designated or indicated in
accordance with the Rules prescribed for determining the type
species solely on the basis of the original publication.

Orthography and Gender of Names and Epithefs

RULE 27. The original spelling of a name or epithet must be retained, except
typographic or orthographic errors. When two or more generic names, or
two or more specific epithets in the same genus, are so similar as to cause
confusion, they are treated as orthographic variants of the same name or
epithet.

NOTE 1. The phrase "original spelling" as used in this rule means the spel-
ling employed when the name of the taxon was validly published. U does
not refer to the use of an initial capital or small letter. Adjectives used as
specific or subspecific epithets change gender endings when necessary to
agree with the gender of the generic name.

NOTE 2. The use of a wrong (or an alternative) connecting vowel or vowels
(or the omission of a connecting vowel) in a generic or subgeneric name or
in a specific or a subspecific epithet gives rise to an orthographic variant.

NOTE 3. When there is doubt as to whether the spellings of the names of two
taxa of the same rank or of two epithets are sufficiently alike to be can-
fused, they should be referred to the Judicial Commission for an Opinion.

NOTE 4. An unintentional typographic or orthographic error later corrected
by the author is to be accepted in its corrected form without prejudice as to
original date and validity of publication.

NOTE 5. The spelling of the name of a taxon or an epithet derived from
the Greek but not transliterated by its author into Latin form in accordance
with classic usage (Appendix A) may be corrected as an orthographic error
by the Judicial Commission and placed in the list of preferred spellings
(see Recommendation 27a). The name of the taxon as originally published
becomes an orthographic variant of the corrected name, but without prejudice
to the validity cf publication by the first author. Specific and other epithets
and names of Greek origin differing merely by having Greek and Latin gender
endings respectively are orthographic variants. Epithets having the same
meaning and differing only slightly in form are considered as orthographic
variants. The genitive and adjectival forms of a personal name are, however,
treated as different epithets.

[99]

700 International Code of Nomenclature

Rule 27. Sect. 7. Orthography and Gender of Names

NOTE 6. Names of taxa and specific epithets are often compound words,
sometimes formed from Latin stems, more often from Greek stems, some-
times (less correctly) from stems from two different languages. Such com-
pound words formed from the same stems may differ only in the connecting
vowel.

a. In the formation of compound names of taxa and of specific epithets
from the Latin the preferred connecting vowel is -i-. Such compound
words that differ in spelling only by having the connecting vowel
-i- or -o- are to be regarded as orthographic variants. Names of taxa
which are such orthographic variants and based upon different types
are to be regarded as homonyms. The spelling used for each taxon
or specific epithet should be that used by the author when the com-
pound was validly published.

b. The combining vowel in Greek compounds in which the first component
is a noun (substantive) or adjective is usually -o-. It may be omitted if
the second component begins with a vowel. It is omitted when the first
component ends in -y (glycyphyllus). Another combining vowel may
be used if there is good Greek precedent, e.g., as in Corynebocterlum.
Names of taxa or words used as specific epithets which are Greek
compounds and which differ only in the presence or absence of the
combining vowel -o-, or have another combining vowel, are to be
regarded as orthographic variants. If the names of taxa are based
upon different types they are to be regarded as homonyms. The
correct spelling is that of the name of the taxon or of the specific
epithet used by the author in its initial and valid publication.

c. Compound words derived from two or more different languages
(nomine hybrido) are to be regarded as orthographic variants if they
differ only in the combining vowel. The spelling first used by an author
in the first valid publication should govern. If the two compounds are
names of taxa having different types they are regarded as homonyms.

NOTE 7. The liberty of correcting a name or epithet must be used with re-
serve, especially if the change affects the first syllable, and above all the
first letter of the name or epithet.

NOTE 8. Intentional Latinizations involving changes in orthography of per-
sonal names, particularly those of earlier authors, must be preserved.

Bacteria and Viruses 101

Rule 27. Sect. 7. Orthography and Gender of Names

NOTE 9. Diacritic signs are not used in names or in specific epithets in bac-
teriology. In names or epithets derived from words with such diacritic signs,
the signs must be suppressed and the letters transcribed as follows:

1. ii, 6 and ii become respectively oe, oe, ue. (German)

2. e, e and e become e. (French)

3. 0 (o), ae (ii) and a become oe, ae and aa, respectively. (Scandinavian)
Where diphthongs are not indicated by special type, the diaeresis should be
used where required to show that two consecutive vowels are to be separately
pronounced (are in separate syllables).

NOTE 10. Certain consonants not found in classical Latin, namely w and y,
and the rarely used k, may be employed in bacteriology for names of taxa
and for specific epithets.

ANNOTATIONS
Rule 27. A typographic error is an unintentional misspelling which
is found in a published name of a taxon or in an epithet. It is not
always easy to determine whether a misspelling is unintentional. If
the author later uses the word correctly spelled, the original spelling
is regarded as unintentional, and the corrected spelling as legitimate.

An orthographic error is one resulting from the incorrect trans-
literation from languages having letters or an alphabet differing from
the Latin. For example, De Bary named an organism Bacillus mega-
terium. It has been urged that classic transliteration from the Greek
would give megatherium as the correct spelling. Both spellings have
been used by subsequent authors. However, De Bary himself con-
sistently used the spelling megaterium. An Opinion was issued con-
firming the use of the spelling megaterium. Similarly an Opinion
was issued confirming the spelling lysodeikticus in the species name
Micrococcus lysodeikticus Fleming as consistently used by the author
and generally accepted by bacteriologists, although the correct trans-
literation from the Greek is lysodicticus.

Botanical Code. The rule is succinct: "The original spelling of
a name or epithet must be retained, except that typographic or ortho-
graphic errors should be corrected."

It is noted that the words "original spelling" refer only to the
spelling of the name when validly published; they do not refer to
typography such as initial capital or small letter.

Zoological Rules state that every "Valid Original Spelling" is to
be retained except for 22 emendations approved in Opinions "here-
tofore" and with the further exception that any author may propose
to the Secretary that an emendation of an original spelling in general
use be approved in the interests of stability and universality. The
proposal becomes provisionally efEective upon publication of a notice

702 International Code of Nomenclature

Rule 27. Sect. 7. Orthography and Gender of Names

in tlie Bulletin of Zoological Nomenclature. If no protest is submit-
ted within a period prescribed by the Commission, the former "Valid
Original Spelling" is added to the appropriate Official Index of
Nomina Repicienda for jiriority but not for homonymy.

Rule 27. Note 1. Recommendation 27h states that s]>ecific epithets
should not be capitalized, even when derived from the name of a
person. If an author capitalized the specific epithet in a new species
name, the epithet is not thereby made illegitimate, but it is recom-
mended that subsequent authors use a lower case initial letter.

Since an adjective used as a specific epithet must agree in gender
with the generic name with which it is used to form a species name,
Bacterium aeruginosum Schroeter 1872 when transferred to the genus
Pseudomonas became Pseudomonas aeruginosa (Schroeter) Migula
1900.

Rule 27. Note 2. The generic name Corynebacterium Lehmann and
Neumann 1896 has the combining vowel -e- rather than the con-
ventional -0-. However, this form was used by the Greeks; for ex-
ample, the Greek lexicon contains compounds such as corynephorus.
Corynebacterium conforms to classic usage. Enderlein (1917) pro-
posed the spelling Corynobacterium. This is to be regarded as an
orthographic variant of Corynebacterium. Had Corynobacteriiun
been proposed as a name for a genus having a different type species
it would have been a later homonym of Corynebacterium.

Rule 27. Note 3. See Provision 5 (p. 131) for procedure in requesting an
Opinion. Several Opinions correcting orthographic errors have been
issued by the Jvidicial Commission. Rule 3 and Rule 4 state that the
name of a family shall be formed by the addition of the suffix -aceae
to the stem of the name of the type genus. One can usually identify
the name of the type genus from the name of the family. Many
generic names in bacteriology are compounds with the final com-
ponent -bacter or -bacterium, as in Azotobacter and Corynebacterium.
The combining form (or stem) of the former ends in -r (genitive
-bacteris) . The combining form of the latter ends in -/ (genitive
-bacterii) . However, family names such as Achromobacteriaceae and
Nitrobacteriaceae have been repeatedly proposed, apparently on the
assumption that Nitrobacter and Achromobacter have a stem ending
in -/. The Judicial Commission in an Opinion ruled that the correct
spelling is Nitrobacteraceae and Achromobacteraceae. However Cory-
nebacteriaceae and Chromobacteriaceae are correctly spelled showing
derivation from Corynebacterium and Chrojnobacterium , respectively.

Bacteria and Viruses 103

Rule 27. Sect. 7. Orthography and Gender of Names

Rule 27. Note 4. When typographical or orthographic errors are
discovered in a publication of a new name, it is an act of courtesy
to call the attention of the author to the error so that he niav himself
publish the correction.

Rule 27. Note 5. The specific epithet in the species name Bacillus
kaustophilus Prickett 1928 had classical usage been followed, would
have been spelled caiistophilus. The Greek kappa should have been
transliterated as a c rather than k. No suggested correction has been
proposed in the literature. Rickettsia akari is generally accepted in
the literature although the generic name from the same root is spelled
Acarus.

Streptococcus eiysipelatos and S. erysipelatis are orthographic
variants; they are different transliterations of the same Greek geni-
tive. Latinization as 5. erysipelatis is preferable.

The genitive noun pasteurii and the adjective pasteurianus.
though derived from the same patronymic Pasteur, are neither ortho-
graphic variants nor homonyms.

Some Latin words were spelled alternatively with an initial vowel
or with an h preceding the vowel. This in some cases was due to two
different transliterations of Greek words with an initial spiritiis asper
(transliterated as /?) . Linnaeus used as a specific epithet the Latinized
Greek abrotonum rather than the classical transliteration habroto-
num; for the species Artemisia abrotonum, the Linnaean spelling
should be followed.

Rule 27. Note 7. The name of Pasteur may be Latinized as Pastor.
Both forms have been used in forming names and epithets: Pasteur-
ella Trevisan 1885; Clostridium pasteurianum Winogradsky 1895;
Urobacillus pasteurii Miguel 1889; Saccharobacillus pastorianus Van-
Laer 1892.

Gaillonclle has been Latinized with resultant Gallionella Ehren-
berg 1835. There is also a Gaillonella B017 de St. Vincent 1823.

Rule 27. Note 8. The only diacritic sign permissible in the spelling
of names and epithets in bacteriology is that of the diaeresis (Gr., a
division) which may be placed above the second of two consecutive
vowels to indicate the two vowels are to be separately pronounced
(are in separate syllables) , and not to be regarded as a diphthong.
For example, the specific epithet aeruginosa (as in Pseudomonas
aerugijiosa) is derived from the stem aer- from aes, aeris z= bronze,
copper, in which initial ae- is a diphthong. However, the Greek
a\] = ae is not a diphthong. Many names and epithets in bacteriology
are derived from the Greek i-^p = aer — air; in such words to indi-
cate that the a and e are in two syllables, the sign of the diaeresis may be
used, hence Aerobacter aerogenes (Kruse) Beijerinck but Bacillus
aeris Chester.

704 International Code of Nomenclature

Rule 27. Sect. 7. Orthography and Gender of Names

Several European languages which employ the Latin alphabet
modify certain vowels by use of diacritical marks.

The modern German umlaut may modify the vowels a, o or
u to a, 6 or il, which may alternately be written ae, oe and ue. The
latter alternative must be chosen when German words are Latinized;
Salmonella schottmuelleri, not S. schottmulleri: Schuetzia, not
Scliiltzia; Actinomyces joersteri, not A. forsteri; Pedioplana lineckelii,
not P. hdckelii; Loehnisium, not Lohnisium.

The French accents of the letter e {e, e and e) are not transferred
when a French word is Latinized.

Some Scandinavian languages have diacritic marks for the vowels
o and a. They are 0 or o, ae or a and a which on Latinization become
oe, ae and aa respectively.

An important group of languages employ the Cyrillic alphabet
(Russian, Serbian, Ukranian, Bulgarian, etc.) . The problem of
transliteration is complicated by the fact that in some cases sounds
are represented by the Cyrillic letters which have phonetically no
equivalent in the Roman alphabet. Romanization is effected by the
use of various diacritic marks as well as by letter combinations.
Latinization of words in languages printed in Cyrillic must be pre-
ceded by romanization, then transliteration into Roman letters with
the conventional diacritic marks. Latinization is completed by
elimination of the diacritic marks and the use of appropriate Latin
endings. Obviously the pronunciation of the letters of the Latinized
word will not correspond to those of the original Cyrillic. (For a
discussion of transliteration of Cyrillic for use in nomenclature see
Paclt, Jiri. Taxon 2:159-166. 1953) .

Botanical and Zoological Codes. There are no essential differences
from the Bacteriological Code. Diacritic marks are suppressed in for-
mation of Latinized words.

Bacteria and Viruses 105

Recommendations 27a, 27b, 27c. Orthography and Gender of Names

RECOMMENDATION 27a. When a name or epithet for use in bacteriological
nomenclature is derived from a Greek word, the transliteration to Latin
form should conform to classic usage. Appendix A (q.v.) to this Code may
be used as a guide. If a later author corrects in a valid publication the
spelling of a name or epithet incorrectly transliterated from the Greek, the
correctly spelled word may be regarded as an orthographic variant of the
name or epithet.

RECOMMENDATION 27b. When it is necessary to choose between words
which have been validly published as names of taxa or as specific or sub-
specific epithets and which differ in spelling only because of faulty or
alternative transliterations of Greek to Latin, or because of alternative
transfer of endings denoting gender or oblique cases (particularly the geni-
tive) from Greek to Latin, it is recommended that choices be governed by
the provisions of Appendix A to this Code, (q.v.).

RECOMMENDATION 27c. For scientific names of taxa it is advisable to use
another font from that used for the remainder of the text, or to space the
letters, or to use some similar device appropriate to the text.
Typewritten scientific names should be underlined.

ANNOTATIONS

Recommejidation 27c. Generally in biology it is customary to print
names of genera and species using a different type from that used
in the body of the text. The most common device is the use of italics
[Bacillus siihtilis) . Occasionally bold face type is used (Bacillus
subtilis). Another device employed is to increase the space between
letters (Bacillus subtilis); this is common in Ger-
man publications.

The Zoological Rules recommend the use of some type other than
that used for the text. However, Follett (1955) in a footnote sug-
gests that the zoological Recommendation "should expressly exclude
the name of any suprageneric category." Many authors in zoology use
italics only for names of species and subspecies, not for names of
families, orders, etc.

The International Code of Botanical Nomenclature adopted by
the Eighth International Botanical Congress, Paris 1954 (published
1956) , uniformly uses "another font than that used for the remainder
of the text," for the names of all taxa, not only for names of species,
but also for the names of genera and all suprageneric taxa. In some
paragraph headings the names are in capitals, in all other places the
names of taxa are in italics. However, in Articles 73 and 74 dealing
with "Orthography of Names and Epithets," there are no directives or
recommendations for such use.

In practice, some botanists and botanical journals use italics (or
other distinctive type) for names of species but roman type for names
of higher groups.

106 International Code of Nomenclature

Recommendation 27c. Orthography and Gender of Names

A pamphlet published for the Royal Society, "General Notes on
the Preparation of Scientific Papers" (1950 p. 16) , under the heading
"Botany" makes the following statement:

"The scientific names of plants are Latin binominals (binomials)
and are printed in italics. The scientific names of higher groups
(orders, classes and families) are printed in ordinary roman type
with a capital if in Latin form."

Bacteria and Viruses 707

Recommendation 276. Orthography and Gender of Names

RECOMMENDATION 27d. When a new name for a genus or subgenus is
taken from the name of a person it should be formed in the following man-
ner:

(1) When the name of the person ends in a vowel or y, the letter a is added,
except when the name of the person ends in o, when ea is added.

(2) When the name of a person ends in a consonant the letters ia are
added, except when the name ends in er when a is added.

(3) When the name of a person has been Latinized and ends in us, this
termination is dropped before adding the sufFix.

(4) The name of a bacterial genus or subgenus may be formed from the
name of a person by the addition of a diminutive ending.

(5) The syllables which are not modified by these endings retain their
original spelling, even with the consonants k and w or with groupings
of vowels or consonants not used in classical Latin.

ANNOTATIONS

Recommendation 27d. Relatively few generic names of bacteria have
been taken from names of persons. Of the approximately 168 bac-
terial generic names recognized in the 7th edition of Bergey's Manual
of Determinative Bacteriology, 22 are from patronymics; of the 18
generic names of rickettsiae, 10 are thus derived.

Recommendation 27 d (1) . Among the generic names of bacteria
derived from names of persons ending in a vowel are Gaffkya, Serraiia,
Noguchia, Nevskia, Beggiatoa, Cowdria and Beneckea. No example
has been found in which the name of a bacterial genus is derived
from a person's name ending in a by the addition of ea. In botany
Collaea is taken from Colla. The generic names thus formed are
feminine nouns. The botanical recommendation is essentially the same.
The Zoological Code recommends that generic names derived from the
names of persons ending in the vowels e, i, o, u or y take the ending
-us, -a or -um. However, in zoology it is relatively rare for an ending
other than -a to be proposed. Of seven examples of such names listed by
Follett all end in -a, all are feminine nouns. The option to create
neuter or masculine nouns is rarely exercised. In zoology, personal
names ending in -a add -ia (in contrast to the botanical and bac-
teriological -ea) in formation of generic names.

Recommendation 27 d (2) . Examples: Escherichia, Erwinia, Dono-
vania, Kiirthia, Pastenria, Nocardia, Borrelia, Rickettsia. The recom-
menation that -a be added to names ending in -er has been com-
monly ignored, and -ia has been added, as in Neisseria and Listeria.
In Zinssera the recommendation has been followed.

The Botanical Code includes the same Recommendation.

The Zoological Rules recommend that personal names ending in
a consonant take the ending -ius, -ia or -iiim, making masculine,
feminine or neuter generic names, as Selysius, Lamarckia. Here again
the alternative of forming a feminine noun is usually chosen.

108 International Code of Nomenclature

Recommendation 27d. Orthography and Gender of Names

Recommendation 27 d (3) . In the earlier writings in biology it was
common practice to Latinize the names of persons. For example: Carl
von Linne usually Latinized his name to Carolus Linnaeus. The
plant generic name Linnnea was formed by dropping the ending -^is
and adding -a.

Recommendation 27 d (4) . Generic names are frequently taken from
the name of a person by adding diminutive and sometimes other
endings.

In biology many suffixes have been used to latinize personal names
for use as generic names. Kuntze {Revisio genernm plantariim 1781)
lists 20 "pure suffixes" (reine Suffixe) from botanical writings. The
use of a diminutive ending is frequent in bacteriology. The following
have been used:

-ella, -iella. Commonly -ella is used as a diminutive suffix to per-
sonal names that do not end in a sibilant: Salmonella from
Salmon, Shigella from Shiga, Pasteurella from Pasteur, Borde-
tella from Bordet, VeiUonella from Veillon, Bartonella from
Barton, Graha?nella from Graham; an exception is Naumanni-
ella from Naumann. Sometimes -iella is added to names ending
in -s or -x: Klebsiella from Klebs; Simonsiella from Simons;
Coxiella from Cox; Rickettsiella from Ricketts; an exception is
Moraxella from Morax. In some cases a consonant is inserted for
euphony: Miy a gate an ell a from Miyagawa. Or a final vowel is
dropped: Brucella from Bruce.
-illus, -ilia, 'illnm. The first of these Latin diminutive endings
has been used by Heller in coining more than twenty names
for genera carved from the genus Clostridium as: Henrilliis
from Henry; Novilhts from Novy; Macintoshillus from Macin-
tosh.
-iota, the name of the smallest letter of the Greek alphabet, has
been used in forming the generic name Colesiota.
Recommendation 27 d (5) . There was no Latin letter lo, but words
containing the letter w are Latinized with retention of the iv as Wol-
bachia from Wolbach. The letter k was used in old Latin, but in
classical Latin was replaced by c. Personal names from languages
other than Latin or Greek retain the k when Latinized, as Kurthia
from Kurth. The letter y was introduced into the Latin alphabet
for the correct transliteration of the Greek letter upsiloji, for which
there was no exact Latin equivalent, and is recognized in bacterio-
logical Latin, as in Wolhynia from Wolhyn.

Bacteria and Viruses 109

Recommendation 27e. Orthography and Gender of Names

RECOMMENDATION 27e. A new specific or subspecific (varietal) epithet
taken from the name of a man may assume either a substantival or an ad-
jectival form.

When the epithet is a substantive, the modernized Latin genitive is formed
in the following manner:

(1) When the name of a person ends in a vowel or -y, the letter -i is
added.

(2) When the name ends in a consonant, the letters -11 are added (thus,
welchii from Welch) except when the name ends in -er, when -i is
added (thus, barker] from Barker).

(3) When the personal name is already Latin or Greek or has been
latinized, the appropriate Latin genitive should be used.

(4) The syllables which are not modified by these endings retain their
original spelling, even with the consonants k and w or with group-
ings of vowels or consonants not used in classical Latin.

When the epithet is an adjective, it is formed by the addition of an ap-
propriate ending.

ANNOTATIONS

Recommendation 27e. A specific epithet taken from the name of a
man is usually placed in the genitive (possessive) case; sometimes an
adjective is formed by use of an appropriate Latin ending.

Names of men that are already Latin or have been Latinized form
the conventional Latin genitive. Carl von Linne Latinized his name
to Linnaeus, with the genitive Linnaei. It is customary to regard names
ending in -a as in the first declension forming the genitive by adding
-e, as Shigae from Shiga, Miyagaivae from Miyagawa.

Names of men in languages other than Latin are first Latinized.
The Romans had several forms of proper names, praenomina, cog-
nomina, names of gentes (clans) . With few exceptions these had -us
as an ending, in most cases the name ended in -ins. In conventional
formation of the genitive the -ins becomes -ii. However, the Latins
noted difficulty in pronouncing genitives of names in which the
-ius is preceded by a vowel. The -ii in such cases was contracted and
became -/; thus the genitive of Gaius is Gai (not Gaii) , of Pom-
peius is Pompei (not Pompeii) ; hence the recommendation that
personal names ending in a vowel other than -a form the genitive
by addition of -/: as Sonnei (not Sonneii) from Sonne, Noguchii
(not Noguchiii) from Noguchi.

Most Latin personal names ending in -ins had a consonant pre-
ceding this ending. Two methods of forming the genitive were com-
monly employed. In perhaps the majority of examples found in
Latin the genitive ending is -ii, but in some it is -i. One finds Plinius,
Plini; Aesculapius, i; Curtius, i. On the other hand scores of names
are recorded which end in -ius and have -ii in the genitive, as
Arcesius, ii; Curtilius, ii; Cuspius, ii; Mallius, ii.

1 10 International Code of Nomenclature

Recommendation 27e. Orthography and Gender of Names

The Bacteriological Code agrees with the Botanical Code in
recommending that in Latinization of non-classic names ending in a
consonant the genitive is to be formed by adding -ii.

The Zoological Rules on the other hand recommend that in all
cases the genitive of non-classic Latin names should be formed by
the addition of -i. The recent (1957) Bradley Draft of the proposed
revision of the Zoological Code accepts the correctness of cither geni-
tive ending -i or -ii.

This difference between the recommendations of the codes is
sometimes confusing. The specific epithets derived from proper
names have frequently been formed by bacteriologists by the addition
of -i. In some cases names of species in different genera may have
specific epithets derived from the name of the same person, one
author used the ending -ii, the other -i. Rule 27 requires that the
original spelling of names or specific epithets should be accepted,
even though not formed in conformity with a recommendation. In
many cases specific epithets in the species names of bacteria have been
changed by a later writer to comply with the Recommendation 27e.

Genitives formed from the personal name of a man by addition
of -ii in the one case and -/ in the other should be regarded as ortho-
graphic variants of the same word. Within the same genus two species
names with specific epithets differing only in the genitive ending
are to be regarded as homonyms.

Some authors "correct" all genitives ending in -i to -// even
though there is no authorization of such change under the rules.

When an epithet in adjectival form is based upon the name of a
man, it is customarily formed by the use of the adjectival endings
-anus or -ianus. When the name ends in a vowel other than -a, the
ending -arms is preferred. If the word ends in -a or -us this ending is
altered to -anus, if the word ends in -ius or -ia or in a consonant the
adjective ends in -ianus. Examples: pasteurianus in Acetobacter pas-
teurianus (from Pasteur) , Acetobacter kuetzijigianus (from Kuetzing) ,
Rhodospirilhun 7nolischianum (from Molisch) , Pseudomonas touel-
liana (from Tonelli) , Xanthomonas hemmiana (from Henuni) .

Bacteria and Viruses 1 1 7

Recommendation 27f. Orthography and Gender of Names

RECOMMENDATION 27f. The same provisions apply to epithets formed from
the names of women. When these have a substantival form they are given
a feminine termination.

ANNOTATIONS

Recommendation 27f. When the name of the woman ends in -a the
genitive is formed by adding -e, as stellae from Stella, marine from
Maria, krzemienieicskae from Krzemieniewska.

The genitives of names of women ending in a vowel other than
-a or that end in a consonant are formed by adding -ae.

Women's names that are in Latin form or that have been latinized
form the genitive in accordance with the rules of Latin grammar, as
Beatricis from Beatrix.

When the specific epithet is an adjective derived from a woman's
name an appropriate adjectival ending is used, usually -amis, -a, -urn,
or -ianus, -a, -um. Inasmuch as the gender of an adjectival specific
epithet agrees with the gender of the generic name, the adjective does
not show definite relationship to the name of a woman. The use of
the genitive substantive may therefore be preferred. The genitive
smithii shows derivation from the name of a man, smithae from the
name of a woman, but smithianus does not indicate whether a man
or a woman is honored.

J 12 International Code of Nomenclature

Recommendations 27g, 27h. Orthography and Gender of Names

RECOMMENDATION 27g. New specific (or other) epithets should be written
in conformity with the original spelling of the words from which they are
derived and in accordance with the rules of Latin and Latinization.
RECOMMENDATION 27h. Specific epithets, even those derived from names
of persons, should not be capitalized.

ANNOTATIONS

Recommendation 27h. During the first several decades following the
introduction of the Linnaean binomial naming of species, it was custo-
mary, particularly in botanical nomenclature, to capitalize all specific
epithets derived from proper names, such as names of persons, places,
mythological characters and genera. Zoologists rather early abandoned
this custom. Later, botanists modified the Recommendation governing
this matter and recognized the right of an author to capitalize or not
as he desired. In bacteriology, as in zoology, the reconmiendation is
that specific epithets begin with a lower case letter no matter what
the origin. This recommendation has some practical significance.
Many abstracting journals and many publications in biology must
use in a single article the scientific names of bacteria, of plants and
of animals. To have different regulations for the capitalization of
specific epithets would be confusing. However, if an author chooses
to capitalize a specific epithet, the species name is accepted, though
those who use the word subsequently may use a lower case initial letter.

Bacteria and Viruses 113

Recommendation 27i. Orthography and Gender of Names

RECOMMENDATION 27i. Names of taxa and specific epithets are often com-
pound words, sometimes formed from Latin stems, more often from Greek
stems, sometimes (less correctly) from stems from two different languages.
Such compound words formed from the same stems may differ only in the
connecting vowels.

(a) In the formation of compound names of taxa and of specific epithets
from the Latin, the preferred connecting vowel is -i-. Such compound
names differing in spelling only by having the connecting vowel -i- or
-o- are to be regarded as orthographic variants. Names of taxa which
are such orthographic variants and based upon different types are
to be regarded as homonyms. The spelling used for the name of each
taxon or for each specific epithet should be that used by the author
when the compound was validly published.

(b) The combining vowel in Greek compounds in which the first com-
ponent is a noun (substantive) or adjective is usually -o-. It may be
omitted if the second component begins with a vowel or when the
first component ends in y, e.g. glycychylus. Another combining vowel
may be used if there is good Greek precedent, e.g., as in Coryne-
bacterium. Names of taxa or words used as specific epithets which are
Greek compounds and differ only in the presence or absence of the
combining vowel -o-, or have another combining vowel, are to be
regarded as orthographic variants. If the names are based upon
different types they are to be regarded as homonyms. The correct
spelling is that used by the author of the name in its initial and valid
publication.

(c) Compound words derived from two or more languages (nomina
hybrida) are to be regarded as orthographic variants if they difFer
only in the combining vowel. If two compounds which are names
of taxa have different nomenclatural types they are to be regarded
as homonyms. The spelling first used by an author in the valid publi-
cation of a name should govern.

ANNOTATIONS

Recommendation 27i (a). The Latins, in contrast to the Greeks,
rarely joined stems to form new words. Wlien stems were so joined,
the connecting vowel was usually -i-, as in atricapillus (black haired) ,
albico7niis (^vhite haired) , aquifolius (with pointed leaf) , in modern
Latin moniliformis (in form of a necklace) . New words were more
commonly formed by use of numerous prefixes and suffixes.

In the modern Latin of biology there are numerous instances in
which the connecting vowel between Latin stems is -o-. This is par-
ticularly true of words in which the first component has to do with
color. The Latins occasionally coined such words as albogiknis
(whitish yellow) in which both components are adjectives, the first
component of the compound having an ad\erbial connotation

114 International Code of Nomenclature

Recommendation 27i. Orthography and Gender of Names

(whitish) . This precedent has been followed in bacteriology in
numerous compound specific eipthets, as witnessed by albidoflavus,
albogilvns, alboniger, albosporeus, atrofaciens, aurogenus, nigro-
maculans, jlavogriseus, jlavovirens, griseoflavus, griseohiteus, roseo-
flmnts, riibropertinctns.

Recommendation 27i (b) . The Greeks formed many compound
words. In consequence, Greek stems have been much used in making
compounds for use as generic names throughout biology. Most Greek
compounds with stems from adjectives and nouns show -o- as the con-
necting vowel. Greek precedent for Corynebacteriuryi is found in Greek
compounds such as corynephorus (club-bearing) . A modern specific
epithet is halmephilus (brine-loving) . Examples of use of -o- are to be
found in many generic names, as Achromobacter, Actinomyces, Blasto-
caulis, and in specific epithets as ochro?nogenes, lipophagus, lepto-
trichoides.

Recommendation 27i (r) . The recommendations advising against
the formation of compound names or epithets from words of two or
more languages, particularly Latin and Greek, have been frequently
ignored. Since such names in their formation do not violate a rule,
they may be legitimate. There are many such words in bacteriology;
some are Latin-Greek, some Greek-Latin, some have -i- as the combin-
ing vowel, some have -o- and some have another vowel.

Latin-Greek, with connecting vowel -o-: Acetobacter, acidopliilus.

albosporeus, jlavochromogenes, Lactococcus.
Latin-Greek with connecting ^'owel -i-: Brevibacterium, Rami-

bacterinm.
Greek-Latin with connecting ^owel -o-: Actinobacilhis, thermo-

fuscus (sic) .
Greek-Latin with connecting vowel -i-: bacteriferus, cristalliferus.
Latin-Greek with connecting ^owel neither -o- nor -i-: Catenabac-

terium.
Greek-Latin with connecting vowel neither -o- nor -i-: gonidia-
formans.

Bacteria and Viruses 115

Recommendations 27\, 27k. Orthography and Gender of Names

RECOMMENDATION 27j. Authors should give the etymology of new generic
names and also of new epithets when the meaning of these is not obvious.
RECOMMENDATION 27k. When it is necessary to choose between words
which have been validly published as names of taxa or as specific epithets
and which differ in spelling only because of faulty or alternative spelling of
Latin words, or because of faulty or alternative transliterations of Greek to
Latin, or because of alternative transfer of endings denoting gender or the
oblique cases, (particularly the genitive) from Greek to Latin, it is recom-
mended that choices be governed by the provisions of Appendix B to this
Code.

ANNOTATIONS

Recommendatioyi 27 j. Compliance with this recommendation might
well tend to reduce the ninnber of nonsense words in bacteriological
nomenclature, particularly nonsense specific epithets, in species names
such as Bacillus alholactis (meaning the Bacillus of white milk in-
stead of the intended white Bacilhis of milk) , or Bacillus thermo-
diastaticus (meaning the "thermally diastatic Bacillus," the intended
meaning probably being the "thermophilic and diastatic Bacillus") .
The student is surprised to find that Streptomyces novaecaesariae
means the "Streptomyces of New Jersey," that noveboracensis refers
to "New York," cantahrigensis to Cambridge and Hajiiia to Copen-
hagen.

Possibly formation of hybrid names (nomina hybrida) such as
Brevibacterium, Acetobacter and Sarcinococcus would be less frequent
if the authors clearly stated that the hybrid word contravened a
nomenclatural recommendation.

716 International Code of Nontenclature

Recommendations 271, 27m, 27n, 27o. Orthography and Gender of Names

RECOMMENDATION 27 I. Words transliterated and Latinized from languages
that do not use the Latin alphabet are to be regarded as orthographic vari-
ants if they differ only in the transliteration. Such orthographic variants used
as names of taxa based upon different types are to be regarded as homo-
nyms.

RECOMMENDATION 27m. Names of taxa and specific epithets difFering only
in the presence or suppression of diacritical marks, transliteration of the
German umlaut or the use of special letters are to be regarded as ortho-
graphic variants; when based upon different types, they should be treated
as homonyms. Words not formed in conformity with Rule 27 Note 9 may be
corrected.

ANNOTATIONS

Recommendation 271. Several different systems of transliterating
Slavic words have been used by German, French, English and other
authors. Different transliterations into the Roman alphabet have
given Metschnikoff, Metchnikov, Metschnikow, all from a Russian
proper name. Specific epithets derived from these varied spellings
are to be regarded as orthographic variants if the differences are in
transliteration only.

The accepted specific epithet lysodeikticiis would be more cor-
rectly Latinized from the Greek as lysodicticus. The two transliterations
would be regarded as orthographic variants of the same word.

RECOMMENDATION 27n. A name or an epithet derived from a non-Latin
or a non-Greek word which does not have an ending conforming to one of
the Latin declensions and which has not been latinized by the addition of a
Latin ending shall be regarded as a non-declinable Latin noun. Two or more
names or specific epithets derived from the same word, one with a Latin
ending, the other not, are orthographic variants, and when the names of taxa
are based on different types they should be treated as homonyms.

RECOMMENDATION 27o. Authors of new names of taxa or of specific or
subspecific epithets derived from the Greek are urged to transliterate words
or stems into Latin form by adherence to the recommendations of Appendix
A.

ANNOTATIONS

Recommendation 27n. Most of the very numerous serotypes of species
of the genus Salmonella have been named by using the unaltered
name of the place of origin (town, city, district) as an epithet with
the name of the genus. In some cases the resultant binomial is used
as the name of the species. The specific epithet in Salmonella lojidon is
to be regarded as a nondeclinable modern Latin word. There is good
Latin precedent for acceptance of such words. One finds many such
words taken over, for example, from the Hebrew in the Biblia Latina
Vnlgata.

Bacteria and Viruses 117

Recommendation 27p. Orthography and Gender of Names

RECOMMENDATION 27p. The name of a genus should be spelled without ab-
breviation the first time it is used with a specific epithet in a publication, (e.g.,
Serratia marcescens) except that in a series of names of species of the same
genus it is customary to abbreviate the name of the genus in all names of the
species after the first. Later use of the name of a species previously cited
usually has the name of the genus abbreviated. The abbreviation used for the
name of a genus in a species name is usually the first letter of the generic name
(e.g., S. marcescens). In publications in which species are listed belonging to
two or more genera which have the same initial letter, care should be used
to abbreviate the names of the genera so as to avoid confusion.

ANNOTATION

Recommendation 27p. The use of the first letter of the name of a
genus as an abbreviation of the name is common practice in biology.
However, some authors seek to avoid confusion when discussing two
or more genera having the same initial letter by using a longer and
more distinctive abbreviation for each. One finds the generic name
Shigella abbreviated as .S'., Sh., and Shig.; Salmonella as S., Sal., and
Salm.; and Streptococcus as S., Str., and Strep. Suggestions have been
made that a standard list of abbreviations should be developed and
adopted. No such list has received any formal approval. There would
arise many difficulties in its formulation. More than twenty generic
names have been proposed in bacteriology which have Thio as a first
component. Even abbreviations as long the following are not dis-
tinctive. Achrom., Actino., Aero., Arthro., Azoto., Bact., Caryo., Cell,
Chloro., Chrom., Haemo., Halo., Lacto., Lepto., Leuco., Methano.,
Micr., Myco., Myxo., Nitro., Nitroso., Oscill, Para., Past., Pelo., Pro.,
Rh., Rhab., Rhodo., Ricketts., Saccharo., Sapr., Sarcin., Sider., Sphaer.,
Spir., Sporo., St., Str., Strepto., Sulfo., Thermo., Thio., Uro., and
Zymo. No proposed or recommended list of such abbreviations has
been widely publicized.

7 78 International Code of Nomenclature

Rule 28. Sect. 7. Orthography and Gender of Names

RULE 28. The gender of generic names is governed by the following regu-
lations:

(a) A Greek or Latin word adopted as a generic name retains its classical
gender. In cases where the classical gender varies, the author has
the right of choice between the alternative genders. In doubtful
cases general usage should be followed.

(b) Generic and subgeneric names which are modern compounds formed
from two or more Greek or Latin words take the gender of the last.
If the ending is altered, the gender is that of the new ending in the
language of origin.

(c) Arbitrarily formed generic names or vernacular names used as
generic names take the gender assigned to them by their authors.
Where the original author has failed to indicate the gender, the next
subsequent author has the right of choice.

ANNOTATIONS

Latin words used as generic names are relatively rare, Latin
compounds are not abinidant. The most common generic names are
Greek compounds; many others are derived from personal names.

Rule 28 (a) . Bacillus, a staff or rod (masculine) and Sarcina, a
packet (feminine) are examples of Latin nouns used without change
as generic names. Vibrio, that which vibrates, is derived from the
Latin and is feminine.

Latinized Greek nouns used as generic names are Bacterium, small
rod or staff, and Clostridium, a small spindle, both neuter.

Rule 28 (b) . A few generic names are compounded from Latin
words, as Lactobacillus, milk rodlet (masculine) , and some hybrid
Greek-Latin as Thiobacillus, sulfur rod (masculine) , and Thiosarcina,
sulfur packet (feminine) .

Throughout biology, including bacteriology, generic names are
commonly Latinized Greek compounds. Examples:

Masculine: Azotobacter nitrogen rod, Actinomyces ray fungus,

Arthrobacter jointed rod, Haemophilus blood lover.
Feminine: Pseudomonas false monad, Crenothrix spring (foun-
tain) hair, Spirochaeta spiral thread, Zoogloea animal glue.
Neuter: Chromobacterium colored rodlet, Mycoderma fungus
skin, Polyangium many vessels, Rhizobium root dweller, Rhodo-
spirillum red spiral, Treponema turning thread.
A few generic names of bacteria do not conform to Recommenda-
tion 5a (4) . They are adjectival in derivation and form. In some
of these cases ithe gender is not indicated by the nominative ending,
e.g., in Alcaligenes and Bacteroides. Custom has fixed these as mascu-
line.

Bacteria and Viruses 1 1 9

Rule 28. Orthography and Gender of Names

The gender of compound generic names having -bacter as the
final component is fixed as masculine by Opinion 3 of the Judicial
Commission (see Appendix C) . A request for an Opinion was based
upon the fact that some authors regarded such names as neuter (per-
haps as a result of confusion with generic names ending in -bac-
terium) , others considered them to be masculine.

Rule 28 (c) . Most Latin names of plants are feminine. This prece-
dent led to the Recommendation both in bacteriology and botany
that new names of genera derived from the names of persons should
be put into feminine form. While the Zoological Code indicates
equal acceptability for generic names of words from personal names
with endings indicating any one of the three genders, nevertheless
in most cases generic names in zoology thus derived have been given
a feminine ending. In bacteriology most generic names of this type
are formed regularly either from personal names, as in Escherichia,
or by addition of a feminine diminutive as in Pasteurella. Even
names not regularly formed, as Ricolesia usually indicate their gender
by the ending.

CHAPTER 4

ProvlsLons for Exceptions to the Rules and

for the Interpretation and

Modification of Rates

CHAPTER 4*

Provisions for Exceptions to the Rules and

for the Interpretation and

Modification of Rules

PROVISION 1

Modification and amendmeiit of Code. This Code can be
amended only by action of the International Committee on Bacterio-
logical Nomenclature and approval of a plenary session of an In-
ternational Congress for Microbiology convened by the International
Association of Microbiological Societies.

PROVISION 2

Lists of nomina co?iser~vanda. To avoid disadvantageous changes
in the nomenclature of the genera by the strict application of the
Rules of Nomenclature, the Rules provide for a list of names which
must be retained as exceptions (nomina conservanda) .

Note 1. This list of conserved names will remain permanently
open for additions. Any proposal of an additional name must be
accompanied by a detailed statement of the case for and against its
conservation. Such proposals must be submitted to the Judicial
Commission (see Provision 4) for study and appropriate action.

Note 2. When a name proposed for conservation has been pro-
visionally approved by the Judicial Commission, bacteriologists are
authorized to retain it pending the decision of the next International
Congress for Microbiology.

Note 3. A conserved name is conserved against all other names
for the taxon, whether these are cited in the corresponding list of
rejected names or not, so long as the taxon concerned is not united
with another taxon bearing a legitimate name. In the event of
union or reunion with another taxon, the earlier of the two com-
peting names is adopted in accordance with Rules 19, 20 and 21.

Note 4. A conserved name is conserved against all earlier homo-
nyms.

* No annotations are included in Chapter 4; the entire chapter is a part of the of-
ficial Code.

[123]

124 International Code of Nomenclature

Provision 3. Provision 4.

PROVISION 3

Lists of nomina rejicienda. To avoid unnecessary confusion in
the nomenclature of bacteria by the strict application of the rules of
nomenclature, the Rules provide a list of names (nomina rejicienda)
which are not to be used, i.e., are to be permanently rejected. This
list includes names which, owing to segregation, are used with dif-
ferent meanings and have become a permanent source of confusion
or error (no77iina nmbigiia) , names where application is uncertain
(jiomina dubia) , and names applied to a group made up of two or
more discordant elements, especially if these elements were errone-
ously supposed to form part of the same individual (fiorniua con-
fiisa) .

Note 1. This list of rejected names will remain permanently open
for additions. Any proposal of an additional name must be accom-
panied by a detailed statement of the case for and against its re-
jection. Such proposals must be submitted to the Judicial Commis-
sion of the Nomenclature Committee for study and appropriate ac-
tion. When a name proposed for rejection has been provisionally
rejected by the Judicial Commission, bacteriologists are authorized to
reject it pending the decision of the next International Congiess for
Microbiology.

Note 2. A rejected name may not be later introduced legitimately
into bacteriological literature, except that nomina dubia may be re-
moved from the list upon submission of evidence of correct status and
by action by the Judicial Commission.

PROVISION 4

Permanent Interyiational Committee on Bacteriological Nomen-
clature. A Permanent International Committee on Bacteriological
Nomenclature has been established by the International Association
of Microbiological Societies in Congress. This Committee is so consti-
tuted that wherever practicable each country is represented by at
least one member and no country by more than five. Nominations
for membership are made preferably by one or more of the micro-
biological societies in each country. If nominations are not made by
the microbiological societies of a country, nominations may be made
by members of the Nomenclature Committee. Nominations should
be made in writing to one of the Permanent Secretaries of the Com-
mittee. Acceptance will be indicated by the Secretaries, and the
nominees given temporary membership in the Committee pending
action of the Committee and of the next International Congress,
when the election will be confirmed.

The several societies and Committee members making nomina-
tions are urged to nominate persons representing all branches of

Bacteria and Viruses 125

Provision 4 cont.

microbiology, so that the International Committee on Bacteriological
Nomenclature may have broad representation of all microbiological
interests in nomenclature. The International Committee on Bac-
teriological Nomenclature through its Permanent Secretaries shall
submit a list of its members to the last Plenary Session of each In-
ternational Congress for confirmation.

At least one year before each Congress, each national society shall
inform the Permanent Secretaries of any recommendations relative
to changes in its representation.

Recognition of Alternates. If a member of the International Com-
mittee on Bacteriological Nomenclature cannot attend the meetings
of the Committee, an alternate having all of the rights of an ap-
pointed member, except in the election of officers and commissioners,
will be chosen in accordance with the following provisions:

(a) The national microbiological society which the member
represents shall have the right to nominate an alternate.

(b) If no nomination is made by the national society, the mem-
ber himself shall have the right to nominate an alternate.

(c) If neither the national society nor the member nominates an
alternate, the Chairman of the Committee with the Perma-
nent Secretaries may nominate.

All nominations for alternates by national societies, or by mem-
bers shall be in writing and should be in the hands of one of the
Permanent secretaries before the first meeting of the Committee.

All nominations for alternates shall be presented at the first
meeting of the Committee, and nominees elected by vote of the
Committee will serve as alternates.

Finictions of the Internatio7ial Committee. The International
Committee on Bacteriological Nomenclature has the following func-
tions:

(1) To hold meetings at the time of and as a part of the sessions
of each Congress of the International Association of Micro-
biological Societies.

(2) To consider and pass upon all recommendations made by
the Judicial Commission relative to the formulation, the
modification or the amendment of the International Code of
Nomenclature of Bacteria and Viruses, or relating to the
nomenclature of other groups of microbiological concern. In
cases in which the approval of the Congress is also necessary the
Committee will make the appropriate recommendations.
Proposals for amendment of the Bacteriological Code shall
be submitted to one of the Permanent Secretaries at least one
year before the next International Congress.

(3) To consider all Opinions issued by the Judicial Commission.
Such Opinions become final unless rejected by majority vote
at the meeting of the Committee next following the date on

126 International Code of Nomenclature

Provision 4 cont.

which the Opinion was published or submitted to the Com-
mittee.

(4) To authorize or appoint special subcommittees of experts
to consider and report on the classification of special groups
of microorganisms and other phases of their taxonomy, to
sponsor one or more sessions of a Section on Taxonomy and
Classification at each International Congress at which these
reports of subcommittees may be presented and discussed;
to receive and recommend for publication the reports of these
subcommittees, and to refer to the Judicial Commission any
special problems that have been raised or recommendations
that have been made respecting nomenclature.

(5) To elect from the membership of the Committee the members
of the Judicial Commission as vacancies may occur and to
replace the members of the several classes as their terms ex-
pire.

(6) Whenever a vacancy occurs, to nominate a Permanent Secre-
tary to the next Plenary Session.

(7) To appoint as Life Members of the International Commit-
tee on Bacteriological Nomenclature individuals who have
rendered distinguished service to bacteriological nomenclature
and taxonomy. Such Life Members shall be regarded as
members-at-large and not as representing the microbiologists
of any nation.

Election of Chairman and Vice-chairman. At a meeting held at
each International Congress, the Committee shall elect a Chairman
and a Vice-Chairman, who shall hold office until the close of the next
succeeding Congress.

A. The Duties of the Chairman of the Committee shall be:

(1) To preside at all meetings of the Committee.

(2) To collaborate with the Permanent Secretaries in the
preparation of the Agenda for the meetings of the Com-
mittee.

(3) To appoint members of such subcommittees as may be
approved by the Committee and not otherwise provided
for.

(4) To assume such other duties as may be requested by the
Committee.

B. The Duties of the Vice-Chairman of the Committee shall be:

(1) To preside at the meetings of the Committee in the ab-
sence of the Chair man.

(2) To assume such other duties as may be requested by the
Committee.

Election of Two Permane7it Secretaries. Two Permanent Secre-
taries are chosen by the Committee and confirmed by the Plenary

Bacteria and Viruses 127

Provision 4 cont.

Session of the Congress. One of the Permanent Secretaries shall
primarily represent the interests of medical and veterinary bacte-
riology and one the interests of non-medical bacteriology. Vacan-
cies in the position of Permanent Secretary which may occur in the
interim between Congresses of the International Association of Micro-
biological Societies shall be filled by the Judicial Commission, such
appointments to be temporary unless confirmed by the next Congress.
The Permanent Secretaries shall be regarded as members-at-large and
shall not be included in the cjuota of any nation.
The duties of the Permanent Secretaries shall be:

(1) To prepare, in cooperation with the Chairman of the Com-
mittee, the Agenda for all meetings of the Committee.

(2) To serve exofficio as voting members and as joint Secretaries
of the Judicial Commission.

(3) To secure from the several national societies representing
microbiology their nominations for membership, and when
such nominations are in accordance wdth the Provisions of
this Code of Nomenclature, to certify such nominees to
temporary membership, and to prepare a list of all nominations
for membership for presentation to the Committee at its first
regular meeting at each International Congress.

(4) To receive from the Judicial Commission such recommenda-
tions as may require action by the Committee; in the intervals
between International Congresses to duplicate and send such
recommendations to all members of the Committee. If the
members of the Committee are circularized to secure com-
ments and suggestions, to tabulate the information received.
If the members are asked to vote upon any proposal, to tabulate
and announce the result of the ballot and to certify the results
to the Chairman of the Committee and to the Chairman of
the Commission.

(5) With the Chairman of the Judicial Commission, to serve as
members of the Editorial Board to edit and prepare for final
publication all Opinions issued by the Judicial Commission,
also the International Code of Nomenclature of the Bacteria
and Viruses and its amendments, and all other publications
authorized by or sponsored by the Committee or by the
Judicial Commission.

128 Infernational Code of Nomenclature

Provision 4 cont.

(6) To keep the minutes of the Committee and of the Judicial
Commission and submit them for publication in the Pro-
ceedings of the appropriate International Congress and in the
International Bulletin of Bacteriological Nomenclature and
Taxonomy.

(7) To present to the Plenary Session of the Congress all actions
of the Judicial Commission or the International Committee
requiring approval of said session.

(8) One or the other of the Permanent Secretaries shall be desig-
nated a member of each subcommittee authorized or ap-
pointed by the International Committee, and shall be re-
sponsible only for advising the specialist subcommittees on
the interpretation of the International Code of Nomencla-
ture of the Bacteria and Viruses.

(9) With the Chairman to authorize the formation of new sub-
committees to consider the taxonomy and classification of
special groups of microorganisms, such subcommittees to have
provisional status only until such time as they may be duly
authorized by action of the International Committee on Bac-
teriological Nomenclature. Proposals for the formation of
such subcommittees may be made by an individual or by a
group of individuals to one of the Permanent Secretaries.
Such a request should be accompanied by a statement of ithe
proposed duties of the subcommittee and a list of the mem-
bers proposed.

Functions of Taxonomic Subcommittees. Taxonomic subcom-
mittees of experts authorized and appointed by the International
Committee on Bacteriological Nomenclature shall work under the
following rules:

(1) The chairman and secretary shall be elected by members of
the subcommittee and shall hold office for six years. After
holding office they shall be eligible for re-election.

(2) Ordinary members of the subcommittee shall be appointed
by the International Committee. Between Congresses, the
chairman and secretary, acting in agreement, may co-opt as
temporary members specialists who can contribute sub-
stantially to the work of the subcommittee. At the next Con-
gress the names of temporary members shall be submitted
to the International Committee, which shall determine
whether they shall become ordinary members.

(3) One of the Secretaries of the International Committee shall
be an ex officio member of each subcommittee, but shall not
have voting powers.

(4) Each subcommittee shall meet at each International Con-
gress and shall review its membership. The names of mem-
bers who have retired or have ceased to interest themselves
in the work of the subcommittee, or who have died, shall be
deleted from the membership list. The subcommittee shall,
if it desires, replace the inactive members by more active ex-
perts, but it is not essential to maintain the membership at
a definite number.

Bacteria and Viruses 129

Provision 4 cont.

(5) The chairman and secretary of each subcommittee will be
jointly responsible for the presentation of a report to the
International Committee. This report, if approved by the
International Committee, will be published in the Interna-
tional Bulletin of Bacteriological Nornenclature and Tax-
onomy. The report should be accompanied by an up-to-date
list of names and addresses of members of the subcommittee.

(6) A subcommittee may create one or more subgroups to study
particular problems. Each subgroup shall have a chairman
appointed by the subcommittee. At least two-thirds of the
members of each subgroup shall be appointed by the subcom-
mittee, but need not themselves be members of the subcom-
mittee; up to one-third may be co-opted as experts, at the
invitation of the subgroup's chairman.

(7) When votes are called for in subcommittees or subgroups, a
simple majority shall suffice; in the event of a tie, the chair-
man shall have a second or casting vote. If members wish to
record a unanimous decision, a fresh vote may be called for.
Votes shall be recorded by a show of hands unless a secret
ballot is demanded by at least one-quarter of the members
present.

(8) Members who cannot attend meetings of subcommittees may
send alternates to act for them. An alternate should be pro-
vided by the member with a written authority which will
entitle him to record a vote on the members behalf. Expert
observers may be introduced by members, but do not have
voting rights.

(9) Between Congresses, the work of the subcommittees will be
conducted by correspondence; when votes are called for, each
ballot paper will be signed by the member; unsigned papers
will not be counted.

130 International Code of Nomenclature

Provision 5.

PROVISION 5

Authorization of a Judicial Commission. A Judicial Commission
is authorized; it shall consist of fourteen members, twelve elected by
the members of the International Committee on Bacteriological
Nomenclature from its membership and the two Permanent Secre-
taries. The Commissioners are elected to serve in three classes of
four Commissioners each, one class retiring at the close of the meet-
ing of each International Congress of the International Association
of Microbiological Societies. In case of the resignation or death of a
Commissioner during the interval between Congresses, the vacancy
may be filled by letter ballot of the members of the International
Committee.

If a Commissioner cannot attend the meetings of the Judicial
Commission, an alternate having all the rights of a Commissioner
except in the election of officers will be chosen in accordance with the
following provisions:

(a) The Commissioner himself shall have the right to nominate
an alternate.

(b) If no nomination is made by the Commissioner, the Judicial
Commission will elect an alternate from members of the
Committee on Bacteriological Nomenclature attending the
Congress.

All nominations shall be in writing and in the hands of the Chair-
man of the Judicial Commission before the first meeting of the Com-
mission. All nominations for alternates shall be presented at the
first meeting of the Commission and nominees elected by vote of the
Commission will serve as alternates.

One of the Commissioners shall be chosen as Chairman and one
as Vice-chairman by vote of the Judicial Commission. The Chairman
and Vice-chairman shall hold office during their unexpired terms as
Commissioners.

Duties of the Chairman of the Judicial Commission. The Chair-
man of the Judicial Commission shall have the following duties:

(1) To preside at meetings of the Judicial Commission.

(2) To prepare, with the collaboration of the Permanent Secre-
taries of the Committee, the Agenda for meetings of the
Judicial Commission.

(3) To appoint such committees as are authorized by the Ju-
dicial Commission but whose appointment has not been
otherwise provided for.

(4) To serve as Chairman of the Editorial Board with the two
Permanent Secretaries. This Editorial Board is charged
with the responsibility of editing and arranging for the
publication of the Opinions issued by the Judicial Com-
mission, the International Code of Nomenclature of Bacteria
and Viruses with amendments, and all other publications

Bacteria and Viruses 737

Provision 5 cont.

issued or sponsored by the Judicial Commission or the Inter-
national Committee on Bacteriological Nomenclature.

(5) With the cooperation of the Permanent Secretaries, to
formulate tentative Opinions and proposals either for dis-
cussion or for vote by the members of the Judicial Com-
mission, and to receive and codify or tabulate the results.

(6) To maintain a list of all requests for Opinions or for as-
sistance in nomenclatural problems which come to the Com-
mission and to expedite the formulation and publication
of Opinions of the Judicial Commission.

(7) With the cooperation of the Permanent Secretaries to put
in form for consideration requests or suggestions for emenda-
tion of the International Bacteriological Code of Nomen-
clature of the Bacteria and Viruses, to circulate such requests
or formulations to the Commissioners, to call for votes, to
tabulate them and to make the appropriate recommendations
to the International Committee on Bacteriological Nomen-
clature.

(8) To transmit, through the Permanent Secretaries, to the In-
ternational Committee the Opinions as issued by the Judicial
Commission, and all recommendations and actions of the
Judicial Commission requiring the consideration or the ap-
proval of the International Committee.

(9) To represent the Judicial Commission on such International
Committees, Boards or Commissions as may be organized
to consider cooperation in biology in the solution of com-
mon problems of nomenclature and taxonomy, particularly
to work with other similar Commissions or Executive Com-
mittees organized for action on problems on nomenclature
in botany and in zoology.

(10) To undertake such other duties as may from time to time

be requested by the Judicial Commission.
Functions of the Judicial Coynmission. The Judicial Commission
has the following functions:

(1) To hold such regular sessions as may be necessary for the
transaction of all business which should come before it at
the time of the sessions of each Congress of the International
Association of Microbiological Societies.

(2) To consider all requests for Opinions relative to the inter-
pretation of the Principles, Rules, Recommendations and
Provisions of the International Code of Nomenclature of
the Bacteria and Viruses where applications are doubtful.
A request for an Opinion should be accompanied by a fully

732 International Code of Nomenclature

Provision 5 cont.

documented statement of the relevant facts; when a request
is not supported by adequate evidence it shall be returned to
the author for revision. When an Opinion is challenged the
basis of the challenge shall be stated and supported by a
documented statement of the relevant facts.

In suitable cases an Opinion shall be prepared which,
if approved by eight or more Conunissioners, shall be issued
and published, becoming thereby available for the guidance
of bacteriologists. All Opinions are reported to the Inter-
national Committee on Bacteriological Nomenclature and,
unless rescinded by majority vote of this Committee, such
Opinions become final.

(3) To consider each proposal for amendment of the Inter-
national Bacteriological Code of Nomenclature and to
formulate a recommendation for such amendment. When
approved by eight or more Commissioners, it shall be sub-
mitted to the members of the International Committee on
Bacteriological Nomenclature and shall be considered as
tentatively approved when it has been accepted by seventy
per cent of the members voting. It shall be submitted for
final approval to the next Plenary Session of an Interna-
tional Congress.

(4) To exercise the plenary powers conferred upon the Com-
mission by the International Bacteriological Code of Nomen-
clature and by the Fourth Congress of the International As-
sociation of Microbiologists to suspend the rules as applied
to any given case, where, in its judgment, the strict appli-
cation of the Rules will clearly result in greater confusion
than uniformity. If the suspension of the rules is by unani-
mous vote of the Commissioners, the decision is final, except
that an appeal may be taken to the International Commit-
tee on Bacteriological Nomenclature, which may rescind the
action by majority vote of those voting. If the suspension of
the rules is initially approved by nine or more Commis-
sioners, but the vote is not unanimous, the resolution for
suspension shall be submitted to the members of the In-
ternational Committee on Bacteriological Nomenclature
and will be validated by an affirmative vote of seventy per
cent of those voting.

(5) To establish lists of the names of taxonomic groups (taxa)
that are to be conserved (nomina conservanda) on the basis
of Opinions issued relative to the status of such names.
There shall be included a list of conserved generic names

{nomina generica conservanda) together with a list of the

Bacteria and Viruses 733

Provision 5 cont.

generic names against which each name is conserved (nomina
generica rejicienda) , also a list of conserved specific epithets
(epitheta specifica conseiiianda) , together with a list of tJie
specific epithets against which they are conserved {epitheta
specifica rejicienda) , also a list of conserved names of taxo-
nomic groups (taxa) of higher ranks than the genus (nomina
consen'anda ordimnn, janiiliariim, etc.) .

(6) To prepare and publish lists of types which have been fixed
through issuance of Opinions of the Judicial Commission.
These may include lists of type specimens, cultures, etc. for
species or subdivisions of species, type species for genera,
type genera for higher taxonomic groups (taxa) . Type
cultures of species and subdivisions of species may be desig-
nated by the number of the strain or other symbol under
which they are canied in the type culture collection.

(7) To prepare and publish lists of names of genera of bacteria
that have been validly published and, if found advisable,
lists of the generic names of other groups in which micro-
biologists are interested. Such lists would be designed to
assist authors publishing new names and combinations to
avoid proposing illegitimate later homonyms.

(8) To prepare and publish a list of publications in the field
of bacteriology in which names proposed shall be regarded
as not validly published and having no standing in bac-
teriological nomenclature.

(9) To report to the International Committee on Bacteriological
Nomenclature at its first meeting at each International
Congress the names of all Commissioners whose terms of
service expire at the close of the Congress and a list of other
vacancies in the membership of the Commission, all of
which should be filled by election by the Committee.

(10) To prepare Opinions, when requested, relative to the nomen-
clatural status of microorganisms studied by microbiological
techniques, but not classed with the bacteria or the viruses,
as for example, the yeasts, molds, algae and protozoa. How-
ever, Opinions shall not be issued with reference to organ-
isms in these groups until they have been approved by the
Commission or Executive Committee charged with the in-
terpretation of the appropriate code of nomenclature
(zoological or botanical) .

(11) Through its Chairman and with the collaboration of the
Permanent Secretaries to cooperate with other Commissions
or similar bodies appointed by the International Botanical
and Zoological Congresses to consider problems of nomen-
clature.

734 International Code of Nomenclature

Provision 5 cont.

(12) Upon approval of the International Committee, to establish
such Trusts or enter into such agreements as may be ad-
visable for the auditing and administration of funds which
may be designated for the payment of the necessary oper-
ating expenses of the Judicial Commission and Committee,
whether such funds originate from grants-in-aid, gifts, royal-
ties, or moneys received from the sale of publications or
from other sources.

(13) To request from appropriate international agencies of the
United Nations, or from other organizations or foundations,
grants-in-aid for the payment of the necessary expenses of
the work of the Judicial Commission and of the Committee.

(14) To sponsor or establish such publications, bulletins, or
journals, as may be found necessary adequately to present
requests for Opinions from the Judicial Commission, to
present discussions of these requests, and to give the steps
from the inception of the study to the final Opinion, sus-
pension of a rule or allocation to lists of nojniyia conservanda
or of nornina rejicienda.

APPENDIX A

Transliteration of Greek Words for Use
in Nomenclature in Bacteriology

APPENDIX A

Transliteration of Greek Words for Use
in Nomenclature in Bacteriology

Authors desiring to vise Greek words in the formation of epithets and
names to be introduced into bacteriological nomenclature are advised to
transliterate such words into Latin form in conformity with recognized
usage as follows:

Greek A, a (alpha) Latin A, a

clktIs olktIvos - actis, actinis

The Greek a is the first letter of two diphthongs.

at - ae. alfxa, alfxaros - haema, haematis.

Rarely and less correctly transliterated as ai as in
oipaXpa - sphaira, whence Sphaira, a protozoan ge-
neric name. Occasionally at does not constitute a
diphthong, as in atarup - aistor and is then trans-
literated ai. Not infrequently at has been incorrect-
ly transliterated as e, as in Hemophilus.

au - au av^o) - auxo

a at - ha and hae, respectively, as in aXs, aXos - hals,
halis
alfxa - haema

Greek B, /3 (beta) Latin B, b

^aKT-qpiov ' bacterium

Greek F, 7 (gamma) Latin G, g

ya\a yaXaKTos - gala, galactis

When 7 precedes 7, k, ^, x. it is transliterated as n.

77 - ng ayyeiov - angium

7K - nc kyKtipoKos - encephalus

7^ - nx aifly^ - sphinx

7X - nch avyxvais - synchysis

[137]

Actinomyces
haematoides

auxinophilus

halophilus
Haemophilus

Bacterium

Galactococcus

Archangium
encephaloides
Sphinx
synchyseus

138 International Code of Nomenclature

Appendix A. Transliteration of Greek

dLirXov-i - (Jif)lus

Greek A, b (delta) Latin /), d

Greek E, e (epsilon) Latin E, e
evrepov - enlerum

The Greek e is the first letter of two Greek diphthongs,
et - /, dtlcv - Ihium

More rarely et - ^ as '^da - zea
tv - eu kv% - ens
e - he cXkos - helcus

^a}OP - Z"J'ff^

Greek Z, ^ (zeta) Latin ^, z
Greek II, r? (eta) Latin E, e

K-qpn'os - cermiis

Greek nouns ending in t/ are feminine. When trans-
literated they were placed in the first declension
with ending a or f as xatrr? - chaete or chaeia

ij - he ryXtos - heliiis

dpl^ - thrix

Greek 6, d (theta) Latin Th, th
Greek I, l (iota) Latin /, i

LxOi's - ichthys
'lttttos - hippus

Greek K, k (kappa) Latin C, c
KopvPT] - coryne

The Greek k is sometimes incorrectly transliterated
as /:, as ^aKT-qpiov - baklerium.

Greek A, X (lambda) Latin L, I

Greek M, p. (mu) Latin M, m

Greek N, v (nu) Latin N, n

Greek H, ^ (xi) Latin A', x

Greek 0, o (omicron) Latin 0, o
o^aiva - ozaena

The Greek o is the first letter of two diphthongs.
OL - oe olbripa - oedema
ov - TTovs - pus

In the Greek, final -of is usually the ending of a neuter
noun or adjective. Such words were placed in the

XeuKos - leucus
pLKpos - micrus
vtKpbs - necrus
pi'^a - myxo

Diplococcus

Enterococcus

Thioploca
zeae

Eubacterium
helcogenes

^oogloea

Spirochaeta
helianthi

Erysipelothrix

ichthyosmius
hippopotami

Corynehacterium

Leuconostoc

Micrococcus

necrophorus

Myxococcus

ozaenae

oedematiens
Rhizopus

Bacteria and Viruses

739

Appendix A. Transliteration of Greek

second Latin declension, with the ending usually
-urn, occasionally -on, as in ayyetov - angium
TrepaiKov - persicon

Final -os is a masculine nominative ending for nouns
and adjectives. Such words were placed in the sec-
ond Latin declension, with the ending usually -us,
occasionally -os, as in (^tXos - philus

Final -os is also found in Greek genitives. Such words
were placed in the third Latin declension with the
ending usually -is, occasionally -os. as in
epufftTreXaros - erysipelatis
h, vos - hys, hyos
6 - ho opfjios - hormus

Greek n, tt (pi) Latin P, p

irXavos - planus

In composition, r followed by a spiritus asper becomes
if -ph.

Greek P, p (rho) Latin R, r

airetpa - spira

Initial p always has the spiritus asper and is transliter-
ated rh, as in pobov - rhodum

In composition, when the second component begins
with p, and the preceding component ends in a
vowel (not a diphthong), the p is doubled, and the
second p retains the spiritus asper.
diappoia - diarrhoea
(vprjKTOS - eurectus

Greek 2, o- s (sigma) Latin S, s

a^paipLKOs - sphaericus

Greek T, r (tau) Latin T, t
r'tTpa - tetra

In composition, r followed by a vowel having a spiritus
asper becomes 6-th.

Greek T, v (upsilon) Latin 7'',y

jui'KTjs, nvK-qros - myces, mycetis

The Greek i; is the second letter of three Greek diph-
thongs.
av - an av^ri - auxe
ev - eu-

ov - u TTovs - pus
V - hy iiSpo - hydro-

Arc hangium
Lycopersicon

hydrophilus

erysipelatis

hyos

Hormiscium

Mycoplana

Spirochaeta
rhodochrous

diarrhoeae
eurectus

sphaericus
tetragenus

Schizomycetes

auxinophilus
Eubacteriales
Rhizopus
Hydrogenomonas

140 International Code of Nomenclature

Appendix A. Transliteration of Greek

Greek *, (p (phi) Latin Ph, ph
(payos - phagus

Greek X, x (chi) Latin Ch, ch
xXa/iuj, x^ttML'^oj - chlarnvs, chlamydis

\p(v8ris - pseudes

ujou - oiim

Greek ^I', xp (psi) Latin Ps, ps
Greek S2, w (omes^a) Latin 0, o

Phagus

Chlamydobacte-
riales

Pseudomonas

Odspora

APPENDIX B

Alternative SpeiUngs of Names and Epithets
in Bacteriology: Orthographic Variants

APPENDIX B

Alternative Spellings of Names and Epithets
in Bacteriology: Orthographic Variants

Alternative spellings (orthographic variants) of names or epithets
found in the literature of microbiology frequently have resulted from
alternative or faulty spelling of Latin words, from faulty or alterna-
tive transliteration of Greek words into Latin form, or from faulty
or alternative endings used to indicate gender or case when Greek
words are transliterated. Authors of new names of taxa and of epi-
thets are urged to make choices in conformity with the following
recommendations:

(a) Alternative spelling of names and epithets taken from the
Latin.

1. Classic Latin precedent should be followed in the forma-
tion of new names of taxa derived from Latin words hav-
ing alternative spellings involving the two diphthongs ae
and oe. The spelling of names of taxa in which the clas-
sic spelling is not observed may be corrected.

2. In formation of new names of taxa from Latin words
spelled classically with an / but alternatively with pli, the
/ spelling is to be preferred. Names of taxa of Latin deri-
vation spelled by the author with ph may be corrected.

(b) Alternative spelling of names and epithets taken from the
Greek.

1. In the formation of a compound name or specific epithet
from a Greek word which increases in the genitive, it is
preferable to use the longer stem unless there is good
Greek precedent for the use of the alternative stem or
combining form; words otherwise similar, derived one
from the short and one from the long form, are not to be
regarded as orthographic variants. If based upon different
types they are not to be regarded as homonyms. However,
when the word constitutes the final component of a generic
name, the longer stem should be used in the formation of
the names of higher taxa; if the shorter stem has been
used the spelling should be corrected.

2. Of two orthographic variants arising from alternative
transliterations of 8t to i or to e, the spelling used by the
author in the first valid publication should be accepted.
If based ujxjn different types they should be regarded as
homonyms.

3. Names of taxa with alternative spellings arising from
transliteration of i] to e or to a are to be regarded as
orthographic variants. If based upon different types, they
are to be treated as homonyms. The preferred spelling is
that used in the first valid publication of the name.

[143]

144 Infernational Code of Nomenclature

Appendix B. Alternative Spellings

4. In the formation of names of taxa and of specific epithets
the Greek diphthong m should be transliterated as ae.
Words in which the diphthong at is transliterated as e
are to be regarded as orthographic variants of those cor-
rectly transliterated. Orthographic variants based upon
different types are to be regarded as homonyms. Names
of taxa in which at has been improperly transliterated
may be corrected.

5. In a Greek compound in which the second or later com-
ponent has p as the initial letter and in which the com-
ponent is preceded by a vowel (but not a diphthong)
the p should be doubled and transliterated as rrh. A
transliteration as r or rh is to be regarded as an ortho-
graphic variant, and if the taxa bearing the names are
based on different types they should be treated as homo-
nyms. Orthographic variants of this kind may be cor-
rected.

(c) Alternative spellings of gender and case endings.

1. Epithets and names of taxa with alternative spellings
arising from the use in Latin of the endings -us and -os
for transliterated masculine nouns and adjectives which
in the Greek end in -os are orthographic variants. Simi-
larly, the alternative spellings arising from the use in Latin
of the endings -tim and -07i for transliterated neuter nouns
and adjectives which in the Greek end in -ov are ortho-
graphic variants. When based upon different types, they
should be regarded as homonyms. When used as the name
of a taxon the correct spelling is that used by the author
in the first valid publication. When used as an adjectival
specific epithet, the Latin endings -us and -um are to be
preferred. Specific epithets may be coiTected to the pre-
ferred spelling.

2. Greek nouns used as specific epithets in the genitive may
have alternative spellings arising from the use in Latin of
the endings -is and -os for transliterated Greek nouns
whose genitive case ending in the Greek is -o?; they are
to be regarded as orthographic variants. The preferred
spelling is -is, and variants may be corrected.

APPENDIX C

OpinLons Relating to Nomenclature
of the Bacteria and Viruses

APPENDIX C

Opinions Relating to Nomenclature
of the Bacteria and Viruses

Foreivord. The first Opinions relating to nomenclature in bac-
teriology resulted from proposals made to the International Com-
mittee on Bacteriological Nomenclature at its meeting in London
held during the sessions of the Second International Congress for
Microbiology in 1936. The Opinions were approved by the Inter-
national Committee and by the plenary session of the Congress.

The International Committee at its meeting held in New York
in 1939 during the sessions of the Third International Congress for
Micro biolog), with the concurrence of the Plenary Session of the
Congress, selected a Judicial Commission with certain functions,
among them:

(a) To issue formal Opinions when asked to interpret rules of
nomenclature in cases in which the application of a rule is
doubtful.

(b) To prepare formal Opinions relative to the status of names
which have been proposed, placing such names when deemed
necessary in special lists, such as lists of nomina coyiservanda,
noynina rejicienda, etc

(c) To develop recommendations for emendations of the Inter-
national Rules for Bacteriological Nomenclature for pre-
sentation to the International Committee.

(d) To prepare formal Opinions relative to types, particularly
types of species and genera, and to develop a list of bacterial
genera which have been proposed with the type species of
each.

[147]

748 International Code of Nomenclature

Appendix C. Opinions A, B, C

These functions were somewhat amplified in the International
Bacteriological Code of Nomenclature approved at the Fourth Inter-
national Congress of Microbiology held in Copenhagen in 1947.

The International Committee (1953) at its meeting in Rome dur-
ing ithe sessions of the Sixth International Congress of Microbiologists
directed that the Opinions voted by this Committee before adoption
of the Bacteriological Code should be designated by letters, those
issued by the Judicial Commission to be numbered.

OPINIONS ISSUED BY THE INTERNATIONAL COMMITTEE ON
BACTERIOLOGICAL NOMENCLATURE IN 1936

OPINION A

Conservation of the generic name Bacillus Cohn 1872, designation of the

type species, and of the type strain of the species
On the proposals of Prof. R. E. Buchanan and Prof. H. J. Conn rela-
tive to the status of the genus Bacillus and its type species Bacillus
subtilis, it was agreed that:

(a) Bacillus Cohn 1872 should be designated as a genus conservondum.

(b) The type species of Bacillus should be designated as Bacillus subtilis
Cohn 1872 emendavit Prazmowski 1880.

(c) The type (or standard) strain should be the Marburg strain.

(d) Cultures of the type (or standard) strain of Bacillus subtilis together
with complete description should be maintained at each of the recog-
nized Type Culture Collections.

(e) The genus Bacillus should be so defined as to exclude bacterial species
which do not produce endospores.

(f) The term Bacillus should be used as a generic name and that it should

be differentiated from the terms "bacillus," "bacille" and "Bazillus"
used as morphological designations.

OPINION B

Generic homonyms in the group Protista

On the proposal by Prof. F. Mesnil relative to generic homonyms, it
was agreed that:

(a) Generic homonyms are not permitted in the group Protista.

(b) It is advisable to avoid homonymy amongst Protista on the one hand,
plant or animals (Metazoa) on the other.

OPINION C

Capitalization of specific epithets derived from names of persons
Prof. R. S. Breed proposed and it was agreed that:

While specific substantive names derived from names of persons
may be written with a capital initial letter, all other specific names
are to be written with a small initial letter.

Bacteria and Viruses 749

Appendix C. Opinions 1-5.

OPINIONS ISSUED BY THE JUDICIAL COMMISSION

OPINION 1

Spelling of the specific epithet in Bacillus megaterium deBary
The spelling megaterium of the specific epithet in Bacillus megaterium deBary
1884 is to be preferred to the spelling megatherium.
(Internatl. Bull. Bact. Nomen. and Tax. 7:35-36. 1951)

OPINION 2

Stems (combining forms) of -bacterium, -bactrum, -bactron, and -bacter
The combining form or stem of the last component of names ending in -bac-
terium is -bacteri, of those ending in -bactrum or -bactron is -bactr, and of
those ending in -bacter is -bacter. Family names derived from such generic
names have respectively the endings -bacteriaceae, -bactraceae and -bac-
teraceae.
(Internatl. Bull. Nomen. and Tax. 7:37-38. 1951)

OPINION 3

Gender of bacterial names that end in -bacter
The names of bacterial genera which end in -bacter should be regarded as
having the masculine gender.

(Internatl. Bull. Bact. Nomen. and Tax. 7:84-85. 1951)

OPINION 4 (Revised)

Rejection of generic name Bacterium Ehrenberg
The bacterial generic name Bacterium Ehrenberg 1828 is to be recognized as
a nomen genericum rejiciendum (rejected generic name). The bacterial family
name Bacteriaceae is to be recognized as a nomen familiae rejiciendum
(rejected family name).
(Internatl. Bull. Bact. Nomen. and Tax. 4:\42. 1954)

OPINION 5

Conservation of the Generic Name

Pseudomonas Migula 1894 and Designation of

Pseudomonas aeruginosa
(Schroeter) Migula 1900 As Type Species

The Editorial Board published a Preliminary Statement (File No.
7) relative to the status of the generic name Pseudomonas Migula
1894 and to the designation of the type species. Three proposals were
submitted to the members of the Judicial Commission.

The first proposal was approved by twelve Commissioners, it was
disapproved by none, and two Commissioners did not vote.

The second proposal was approved by twelve Commissioners, it
was disapproved by none, two Commissioners did not vote.

The third proposal was approved by eleven Commissioners, two
Commissioners did not vote, and one Commissioner stated that in his

150 International Code of Nomenclature

Appendix C. Opinions 5, 6.

opinion the "selection of type species" is made "in accord with
generally accepted rules thus making formal action unnecessary."
The Opinion as approved by the Judicial Commission is as follows:

Opinion 5.

1. The generic name Pseudomonas Migula 1894 is to be conserved and
placed in the list of nomina generica conservanda.

2. The generic name Pseudomonas Migula 1894 is to be associated
with the species designated and described by Migula 1895.

3. The type species of the genus Pseudomonas Migula 1894 is Pseudo-
monas aeruginosa (Schroeter) Migula 1900 Bacterium aeruginosum
Schroeter 1872, Bacillus pyocyaneus Gessard 1882, Pseudomonas
pyocyanea Migula 1895.

REFERENCES

Editorial Board. Status of Pseudomnnns Migula 1894 as a Generic Name

and of Pseudomonas aerus^inosa (Schroeter) Migula 1900 as Type Species

Internatl. Bull. Bad. Nomen. and Tax. / (H :41-42. 1951.
Judicial Commission, Opinion No. 5. Internatl. Bull. Bact. Nomen. and Tax.

2:121-122. 1952.
Migula, W. Ueber ein neues System der Bakterien. Arb. Bakt. Inst. Karlsruhe.

i:237. 1894.
. Schizomycetes (Bacteria, Bacterien) in Engler, \. and K. Praiitl. Die

Naturlichen Pflanzenfamilien. Teil 1. Abt. la. 29. 1895.

. System der Bakterien. 2:884. 1900.

Schroeter, J. Ueber einige durch Bacterien gebildete Pigments, in Cohn, F.,

Beitrage z. Biologic d. Pflanzen. 1 (Heft 2) :126. 1872.
Gessard, C. Sur les Colorations bleue et verte des linges a pausements. C. R.

Acad. Sci. Paris. P^:536. 1882.

OPINION 6

Conservation of the Generic Name Chlorobacterum
Lauterborn 1915 against Chlorobocterium Guillebeau 1890

The currently used bacterial generic name CJilorobacterium Lauter-
born 1915 is a later homonym of ChlorohacterUun Guillebeau 1890.
The latter name has not been used in the literatiue of bacteriology
for more than half a century. A Preliminary Statement (1951) by
the Editorial Board outlining the problem was published, likewise
a series of suggestions as to alternative actions (1953) . At the meet-
ing of the Judicial Commission on September 4, 1953, it was agreed
that the existence of the earlier homonym Chlorobacteriiim Guille-
beau should not constitute a bar to the use of the later homonym
Chlorobacteriiim Lauterborn, and that the name Cholorobacteriiitu
Guillebeau should be rejected.

Opinion 6.

The generic name Chlorobocterium Guillebeau 1890 is placed in the list of

nomina generum rejicienda.

Bacteria and Viruses 151

Appendix C. Opinions 6, 7.

REFERENCES

Editorial Board. Chlorobacterium Guillebeau 1890 vs. Chlovobacterium

Lauterborn 1915, and the status of Chlorobacteriaceae Lauterborn 1913.

Prelim. State. File No. 8. Internatl. Bull. Bact. Nomen. and Tax. 1 (1) :

43-44. 1951.
. Summary of Opinions and Preliminary Statements. Ibid. 3 (2, 3) :

77-78. 1953.'
Guillebeau, Alfred. Studien iiber Milchfehler und Euterentziindungen bei

Rindern und Ziegen. I. Ueber Ursachen der Euterentziindung. Landw.

Jahrb. Schweiz. -/: 27-44. 1890.
Joint Secretaries. Minutes of the Judicial Commission Meetings held in Rome

(1953). Internatl. Bull. Bact. Nomen. and Tax. 3 (4) :149. 1953.
Judicial Commission. Opinion 6. Internatl. Bull. Bact. Nomen. and Tax.

■f (3-4) :143. 1954.
Lauterborn, Robert. Zur Kenntnis einiger sapropelischer Schizomyceten.

Allg. Bot. Zeitschrift. 79:99-100. 1913.
. Die sapropelische Lebewelt. Verhandl. d. Naturhistor. Medizin.

Verein szu Heidelberg i3:425. 1915.

OPINION 7

Nomenclature of the Organism Associated with
Granuloma Venereum

Bier (1953) outlined the problem of determination of the name to
be applied to the organism described by Aragao and Vianna (1913)
as associated with and the probable cause of granuloma venereum.
The synonymy is confused in the literature. The Judicial Commis-
sion, at its meeting of the 5th September, 1953, after a review of the
evidence, approved the following Opinion.

Opinion 7.

The bacterial species names Encapsulatus inguinoJis Bergey et al. 1923,
Klebsiella granulomatis Bergey et al. 1925, Donovania granulomatis Ander-
son, de Monbreun and Goodpasture 1944 are later synonyms of Caiymmato-
bacterium granulomatis Aragao and Vianna 1913.

REFERENCES

Anderson, K., W. A. DeMonbreun and E .W. Goodpasture. An etiologic

consideration of Donovania granulomatis cultivated from granuloma

inguinale (three cases) in embryonic yolk. J. Exp. Med. 57:25-38. 1945.
Aragao, H. de B.. and G. Vianna. Mem. Instituo Oswaldo Cruz. 5:211-238.

1913.
Bergey, D. H. in Bergey's Manual of Determinative Bacteriology, Williams

and Wilkins, Baltimore. 1st Ed. p. 238. 1923. 2nd Ed. p. 264. 1925.
Bier, Otto G. The nomenclature of the bacterium found in granuloma

venereum. Internatl. Bull. Bact. Nomen. and Tax. 3 (1) :20. 1953.
Editorial Board. Summary of Opinions and Preliminary Statements. Ibid.

3 (2,3) :79. 1953.
Joint Secretaries. Minutes of the Judicial Commission Meetings held in Rome

(1953) . Internatl. Bull. Bact. Nomen. and Tax. 3 (4) :150. 1953.
Judicial Commission. Opinion 7. Ibid. 4 (3-4) :144. 1954.

752 Infernational Code of Nomenclature

Appendix C. Opinion 8.

OPINION 8

The Correct Species Name of the Streptococcus
of Bovine Mastitis

Haupt (1952) proposed that the name of the streptococcus of bovine
mastitis be accepted as Streptococcus agalactiae Kitt, although the
binomial given by Kitt was Streptococcus agalactiae contagiosae. Han-
sen (1953) suggested that Kitt's single specific epithet of two words
was changed by Lehmann and Neumann (1896) to a single word,
and that the species name Streptococcus agalactiae Lehmann and
Neumann has become generally recognized and is in common use. A
poll of those working with the bovine mastitis streptococci showed
general agreement that confusion would best be obviated by conser-
vation of the Lehmann and Neumann species name against all syno-
nyms having priority. The Judicial Commission, in its meeting of
5 September, 1953, approved in principle the conservation of the
species name Streptococcus agalactiae Lehmann and Neumann, but
with instructions to the Editorial Board to discuss certain matters
with the proponents of the conservation. It is recognized that
the description of the characteristics of the species as given by
Lehmann and Neumann are probably inaccurate in some particulars
and differ from the descriptions of earlier writers. It is proposed
that a type culture or standard culture of Streptococcus agalactiae
Lehmann and Neumann be selected by a committee of experts, ap-
proved, and adequately described, and that the species be based upon
this type culture rather than on the Lehmann and Neumann de-
scriptions.

Opinion 8.

The species name Streptococcus agalactiae Lehmann and Neumann 1896 is
conserved against all synonyms having priority.

REFERENCES

Editorial Board. Summary of Opinions and Preliminary Statements. Inter-

natl. Bull. Bact. Nomen. and Tax. 5 (2, 3) :79. 1953.
Hansen, P. Arne. A note on the name Streptococcus agalactiae Lehmann and

Neumann. Ibid. 3 (1) :21-23. 1953.
Haupt, Herbert. The correct species name of the streptococcus of bovine

mastitis. Ihid. 2 (3) : 104-106. 1952.
Joint Secretaries. Minutes of the Judicial Commission meetings held at

Rome in connection with the VI International Congress for Microbiology,

Sept. 1953. Ihid. 3 (4) :150. 1953.
Kitt, Th. Bakterienkunde und pathologische Mikrokospie fiir Tieriirzte und

Studierende der Tiermedizin. 2nd Ed., Vienna, 1893.
Judicial Commission. Opinion 8. Ihid. 4 (3-4) :145. 1954.

Bacteria and Viruses 153

Appendix C. Opinion 9.

OPINION 9

Conservation of the Bacterial Generic

Name Gallionella

The species name Gallionella ferruginea Ehrenberg 1838 was applied
by the author to an organism which he erroneously regarded as a
diatom, later shown to belong with the bacteria. A Preliminary
Statement (1952) by the Editorial Board pointed out that the name
Gallionella was an altered spelling (probably an attempt at better
Latinization of the name Gaillonelle) of the earlier generic name
Gaillonella. The ascription of an iron bacterium to a genus of dia-
toms caused Griffith (1853) to propose the new generic name Didymo-
helix for the species D. ferruginea (Ehrenberg) Griffith. The generic
name Didymohelix has not been commonly accepted or used by
bacteriologists, the species name Gallionella ferruginea has almost
universally been used. The name Gallionella for a genus of diatoms
has quite disappeared in the literature of algology.

The botanist's subcommittee on Bacillariophyta (diatoms) pro-
posed that the generic name Gaillonella Bory (1823) be placed on the
list of botanical nomina rejicienda. The reasons for and the impli-
cations of this action were ably analyzed by Ross (1952) . This re-
jection also automatically rejects Gallionella as a generic name of
diatoms. It was suggested that there remained no botanical reason
for any "objection to conserving Gallionella in the sense of a genus
typified by G. ferruginea Ehrenberg."

The Judicial Commission at the meeting 5th September 1953, in
Rome approved the conservation of Gallionella Ehrenberg as a bac-
terial generic name.

Opinion 9.

Gallionella Ehrenberg is placed in the list of conserved names of bacterial
genera (nomina generum conservando) with the type species Gallionella
ferruginea Ehrenberg.

REFERENCES

Bory de St. Vincent. Diet. Classique d'Hist. Nat. 4:Wi. 1823.

Editorial Board. Status of Gallionella Ehrenberg as a generic name in bac-
teriology. Prelim. Statement File No. 49. Int. Bull. Bact. Nomen. and
Tax. 2 (3) :96-103. 1952.

. Summary of Opinions and Preliminary Statements. Ibid. 3 (2, 3) :80-

81. 1953.

Ehrenberg, C. G. Die Infusionsthierchen als voUkommene Organismen. p. 166.
Leipzig. 1838.

Griffith, J. W. On Gallionella ferruginea (Ehrenberg) . Ann. and Mag. Nat.
Hist. Ser. 2. 72:438. 1853.

Joint Secretaries. Minutes of the Judicial Commission meetings held at
Rome in connection with the VI International Congress for Micro-
biology, Sept. 1953. Internatl. Bull. Bact. Nomen. and Tax. 3 (4) :150.
1953.

154 International Code of Nomenclature

Appendix C. Opinions 9—1 1 .

Ross, R. in Nomenclatural Comments. Conservation of GnlUonelln. Ibid.

3 (2, 3): 117-1 19. 1953.
Judicial Commission. Opinion 9. J bid. 4 (3-4) :147. 1954.

OPINION 10

Invalidity of the Bacterial Generic Name Mullerina de
Petschenko 1910 and of the Species Name Mullerina paramecii

De Petschenko (1910) published the name Mullerina for a genus of
bacteria and included one species M. paramecii. In this preliminary
paper he stated that a more extensive description of the organism
would be given later. In a subsequent paper (1911) the names
Drepanospira and Drepanospira muelleri were proposed and used.
An Opinion was asked as to the nomenclatural status of these names
for the organisms described. A preliminary statement was issued by
the Editorial Board (1952) . The following Opinion was approved
by the Judicial Commission at its meeting on 5th September, 1953.

Opinion 10.

The generic name Mullerina de Petschenko 1910 and the species name Mul-
lerina paramecii de Petschenko 1910 were not accepted by the author, hence
were not validly published and are without standing in nomenclature. The
later names Drepanospira de Petschenko 1911 and Drepanospira muelleri
de Petschenko 1911 were validly published and are not later synonyms.

REFERENCES

De Petschenko, Boris. Contributions a I'etude de I'immunite chez les proto-
zoaires. Zentr. f. Bakt. Abt. I. Orig. 56:90-92. 1910.

. Drepanospira millleri n. g., n. sp. parasite des Parameciums; contri-
bution a r^tude de la structure des bact^ries. Arch. f. Protistenkunde
22:248-298. 1911.

Editorial Board. The generic names Miillerina and Drepanospira in bac-
teriology. Preliminary Statement. File No. 52. Internatl. Bull. Bact.
Nomen. and Tax. 2 (1) :9-10. 1952.

. Summary of Opinions and Preliminary Statements. Ibid. 3 (2. 3) :81.

1953.

Joint Secretaries. Minutes of the Judicial Commission Meetings held in Rome
(1953). Ibid. 3 (4) :150. 1953.

Judicial Commission. Opinion 10. Ibid. 4 (3-4) :147. 1954.

OPINION 11

Nomenclature of Species in the Bacterial Genus Shigella

A request for an Opinion as to the correct names of certain species
in the genus Shigella was made by Boyd and Cowan (1951) . It was
supported by the recommendations of the Shigella Commission of
the Enterobacteriaceae Subcommittee (1951). A Preliminary State-
ment was published (1951). The Judicial Commission in its meet-
ing on 5 September 1953, approved the following Opinion.

Bacteria and Viruses 155

Appendix C. Opinion 11.

Opinion 11.

1. Shigella dysenteriae (Shiga) Castellani and Chalmers 1919 was validly

published and is legitimate as the name of the dysentery bacterium de-
scribed by Shiga (1898).

2. The specific epithet flexneri in the species name Shigella flexneri Castellani
and Chalmers 1919 is designated as a conserved specific epithet (epitheton
specificum conservondum) for the species first described as Bacillus dysen-
teriae Flexner 1900.

3. The species name Shigella boydii Ewing 1949 was validly published and
is legitimate. The specific epithet boydii in the species name Shigella
boydii is to be conserved (epitheton specificum conservondum).

4. The species name Shigella sonnei (Levine) Weldin 1927 was validly pub-
lished and is legitimate. The specific epithet sonnei in the species name
Shigella sonnei is to be conserved (epitheton specificum conservondum).

5. A type or standard culture is to be designated by the Enterobacteriaceae
Subcommittee of the International Committee on Bacteriological Nomen-
clature for each of the four species. Such cultures as far as possible shall
be maintained in each of the national Type Culture Collections and in the
International Shigella Center, Chamblee, Georgia, U. S. A.

6. A culture belonging to the species Shigella flexneri, S. boydii or S. sonnei

could be completely identified by appending to the name "serotype "

(with the appropriate designation).

REFERENCES

Bovd. J. S. K. and S. T. Cowan. Shigella species nomenclature and citation

of authority. Request for an Opinion by the Judicial Commission.

Ir.ternatl. Bull. Bact. Nomen. and Tax. 1 (4)' : 147-149. 1951.
Castellani, A. and A. J. Chalmers. Man. Trop. Med. 3rd Ed. p. 9.H5. 1919.
Editorial Board. Shigella species nomenclature and citation ol authority.

Preliminary Statement. File No. 54. Internatl. Bull. Bact. Nomen. and

Tax. / (4) : 158- 16(1. 1951.
. Summary of Opinions and Preliminar\ Statements. Ihid. 3 (2. 3) :82.

1953.
Enterobacteriaceae Subcommittee. Nomenclature and classification of the

dysentery bacilli. Ibid. 1 (4) : 150-157. 1951.
Ewing. William H. Shigella nomenclature. J. Bact. 57:633-638. 1949.
Flexner. Simon. On the etiology of tropical dysentery. Phila. Med. J. 6:414.

1900.
Joint Secretaries. Minutes of the Judicial Commission held at Rome in

connection with the VI International Congress for Microbiology Sept.

1953. Internatl. Bull. Bact. Nomen. and Tax. 5 (4) :150. 1953.
Judicial Commission. Opinion 11. Ibid. 4 (3-4) :148. 1954.
Levine, Max. Some differential characters of the group of dysentery bacilli.

J. Inf. Dis. 27:31. 1920.
Shiga, K. Ueber den Erreger der Dysenterie in Japan. Zent. f. Bakt. Abt. I. 23:

599-699. 1898.
Weldin, John C. The colon-typhoid group of bacteria and related forms.

Relationships and classification. Iowa State Coll. J. Sci. i: 121-197. 1927.

156 International Code of Nomenclature

Appendix C. Opinions 12, 13.

OPINION 12

Conservation of Listeria Pirie 1940 as a
Generic Name in Bacteriology

Listerella Pirie 1927 was proposed as a bacterial generic name. When
the attention of the author was called to the fact that Listerella Pirie
was a later homonym of the generic name of a myxomycete, he (1940)
proposed to replace Listerella by Listeria. This replacement has in
general been accepted by those working in medical bacteriology.
Becker (1951) noted that Listeria Pirie is a later homonym of Listeria
Necker 1790 applied to a genus of flowering plants. Under the rules
of the International Code of Nomenclature of Bacteria and Viruses
Listeria Pirie is illegitimate. However, the suppression of both Lis-
terella and Listeria as bacterial generic names would cause much con-
fusion and resentment.

The generic name Listeria Necker was apparently validly pub-
lished to include certain segregates from the genus Oldenlandia in
the family Ruhiaceae. The name has not been used by botanists
since the time of its introduction (over a period of more than a
century and a half) . The Judicial Commission at its meeting on
September 4, 1953, agreed to conserve Listeria Pirie 1930 as a generic
name in bacteriology and approved the following opinion.

Opinion 12.

Listeria Pirie 1940 (type species Listeria monocytogenes (Murray, Webb,
and Swann) Pirie 1940) shall be placed in the list of conserved names of
bacterial genera (nomine generum conservanda).

REFERENCES

Becker, Elery R. The legitimacy of certain generic homonyms in bacteriology
and protozoology. Internatl. Bull. Bact. Nomen. and Tax. 1 (3) .103-112.
1951.

Editorial Board. Status of Listeria Pirie 1940 as a generic name in bac-
teriology. Preliminary Statement. File 49. Ibid. 3 (2, 3): 109-110. 1953.

. Status of names of bacterial genera that are later homonyms of names

of protozoan genera. Preliminary Statement. Ibid. 5 (2. 3) : 109-1 10. 1953.

Judicial Commission. Opinion 12. Ibid. 4 (3-4): 150. 1954.

Necker, N. J. Elementa botanica. i:206. 1790.

Pirie, J. H. H. A new disease of veld rodents, 'Tiger River Disease." Publ.
South African Inst. Med. Res. 5:163-186. 1927.

. The genus Listerella Pirie. Science 97:383. 1940.

OPINION 13

Conservation and Rejection of Names of Genera
of Bacteria Proposed by Trevisan 1842-1890

Between the )ear 1842 and 1890 Tre\isan proposed and for the most
part validly published at least thirty-three new names for genera and
subgenera of the bacteria. Some of these names have been accepted
and have come into common use in bacteriology, others for one rea-

Bacteria and Viruses 157

Appendix C. Opinion 13.

son or another, have been ignored and not employed. The evaluation
and appropriate disposition of these names from the standpoint of
their legitimacy is an important problem. As an aid to its solution,
Trevisan's "Generi e le Specie delle Batteriacee" Avas photograpically
reproduced (1952). This paper was originally published as a sepa-
rate brochure by the author; copies are rare (none known in
America) . The Editorial Board (1953) discussed the status of the
se\eral generic names proposed by Trevisan, and suggested a series
of draft proposals as to their disposition. The Judicial Commission
was asked to review these proposals and, as far as practicable, to fix
the present status of each of the names. The Commission has placed
the names in four groups. Those that can be adequately identified,
ha\e priority, and are in current use, are placed in the list of generic
names to be conser\'ed. Those that are later synonyms or homonyms
are listed as illegitimate. Those that cannot be identified with rea-
sonable definiteness are placed in the list of rejected generic names.
The status of two names is left indeterminate and requiring more
study. The Judicial Commission at its meeting of 5 September 1953
approved the following Opinion.

Opinion 13.

1. Generic names proposed by Trevisan placed in the list of conserved
generic names (nomina generum conservanda).

Names of Genera and Subgenera Type Species

Beggiaton Trevisan 1842 Beggiatoa alba (Vaucher)

(p. 56) Trevisan 1845

(Oscillaria alba Vaucher 1803)
Klebsiella Trevisan 1885 Klebsiella pneumoniae

(p. 105) (Schroeter) Trevisan 1887

(Bacterium pJieumoniae
croiiposae Zopf 1885)
KurtJiia Trevisan 1885 Kurthia zopfii (Kurth)

(p. 92) Trevisan 1885

(Bacterium zopfii Kurth 1883)

Leptotrichia Trevisan 1879 Leptotrichia buccalis (Robin)

(p. 138) Trevisan 1879. (Leptothrix

buccalis Robin 1853)
Xeisseria Trevisan 1885 Neisseria gonorrhoeae

(p. 105) Trevisan 1885

Pasteurella Trevisan 1887 Pasteurella cholerae-gallinar-

(P- 94) urn Trevisan 1887^

2. Generic names proposed by Trevisan placed in list of rejected generic
names (nomina generum rejiciendo).

Babesia Trevisan 1889 Babesia xanthopyretica Tre\i-

(p. 29) san 1889.

158 International Code of Nomenclature

Appendix C.

Opinion 1 3.

Ii(irlryi()j)sis Trevisaii 1885

(p. m)

Billet in Trevisan 1889
(p. 11)

Cenomesin Trevisan 1889

(p. 1039)
Corvilia Trevisan 1889

(p. 21)

Diaxcid Trevisan 1889

(p. 26)
Eiicornilio Trevisan 1889

(p. 21) (Subgenus)

Euman tcgazzaea Trevisan
1889 (p. 942) (Suboenus)

Eupaciula Trevisan 1889
(p. 23) (Subgenus)

Eiisj)irinu7n Trevisan 1879
(p. 24) (Subgenus)

Leptotrichiella Trevisan
1889 (p. 935) (Subgenus)

Manlegazzaea Trevisan
1889 (p. 137)

Octopsis Trevisan 1885
(p. 102)

Perroncitoa Trevisan 1889
(p. 29)

Pleurosporo Trevisan 1889
(p. 22) (Subgenus)

Pseudospira Trevisan 1889
(p. 23) (Subgenus)

Pseudospirillum Trevisan
1889 (p. 25) (Subgenus)

Bartcriopsis rasmiissenii

Trevisan 1885. (Leptothrix

xnn topyreticns Trevisan
Billetia laminarine (Billet)

Trevisan 1889. {Bacterium

Inniinariae Billet 1888)
Cenomesia albida Tievisan

1889
Cornilia ahei (Fliigge)

Trevisan 1889. (Badllus

alvei Fliigge 1886)
Dicoccia glossophila Trevisan

1889
Cornilia (Eucoruilia) ah'ei

Trevisan 1889. (Bacillus alvei

Cheshire and Cheyne 1885)
Manlegazzaea (Eurnautegaz-

zaea) cienkowskii Trevisan

1879
Pacinia (Eupacinia) putrifica

Trevisan 1889. (Bacillus

putrificus coli Fliigge 1886)
Spirillum (Euspirillum) undula
(Mueller) Ehrenberg 1830.

(Vibrio undula Mueller

1773)
Leptotrichia (Leptotricltiella)

amphibola Trevisan 1889
Man tegazzaea cien k owsk ii

Trevisan 1879
Octopsis cholerae-gallinarum

Trevisan 1885. (Micrococcus

cholerae-gallinarum Zopf

1885)
Perroncitoa scarlatinosa

(Trevisan) Trevisan 1889.

(Micrococcus scarlatinosus

Trevisan 1879)
Cornilia (Pleurospora)

tremula (Koch) Trevisan

1889. (Bacillus tremulus

Koch 1877)
Paciyiia (Pseudospira) cholerae-

asiaticae Trevisan 1885.

(Vibrio cholerae Pacini 1854)
Spirillum (Pseudospirillum)

amphibolum Trevisan 1889

Bacteria and Viruses 759

Appendix C. Opinion 13.

3. Trevisan's generic names which, as later homonyms or synonyms, are
regarded as illegitimate.

BoUingern Trcvisan 1889 BolUngera eqiii (Rivolta)

(p. 26) Trevisan 1889. (Zoogloea

j)ubno7TJs equi Bollinger
1870)
Rasmiissenia Trevisan 1889 Rasmiissenin hxiccalis (Robin)

(p. 930) Trevisan 1889. {Leptothrix

huccalis Robin 1853)
Scliuetzia Trexisan 1889 Schuetzia poelsii Trevisan 1889.

( p. 29) {Streptococcus equi Sand and

Jensen 1888)
Winogradskya Trevisan 1889 Winogradskya ramigera

(p.' 12) (Itzigsohn) Trevisan 1889.

(Zoogloea ramigera Itzigsohn
1867)

4 Trevisan's generic names whose status is indeterminate.

Gafjhya Trevisan 1885 Gaffkya tetragena (Gaffky)

(p. 105) Trevisan 1885. (Micrococcus

tetragenus Gaffky 1883)

Pacinia Trevisan 1885 Pacinia cholerae-asiaticae

(p. 83) Trevisan 1885

REFERENCES

Billet, A. Sur le cycle evolutif et les variations morphologiques d'une nou-
velle bacteriacee marine, Bacterium laminariae. C. R. Acad. Paris. 106:
293-295. 1888.

Bollinger, Otto. Mycosis der Lunge beim Pferde. Virchows Arch. f. path.
Anat.. Berlin. •/5>:583-586. 1870.

Cheshire, F. R., and W. W. Cheyne. The pathogenic history and history un-
der cultivation of a new Bacillus (B. alvei) , the cause of a disease of the
hive bee hitherto known as foul brood. J. Roy. Microscop. Soc. Ser. 2,5s
581-601. 1885.

De Toni. J. B., and V. Trevisan. Schizotnycetaceae Naeg. in Saccardo, P. A.
Syll. Fung. 5:923-1087. 1889.

Editorial Board. Status of generic names of bacteria proposed by Trevisan.
Internad. Bull. Bact. Nomen. and Tax. 3:87-108. 1953.

Ehrenberg, C. G. Beitrage zur Kenntniss der Organisation der Infusorien
und ihrer geographischen Verbreitung besonders in Sibirien. Abhandl. d.
math, und phys. Kl. Akad. Berlin 1-88. 1830.

Fliigge. C. Die Mikroorganismen. Leipzig. 1886.

Gaffky. Georg. Arch. f. "Chirurg. 25:500. 1883.

Itzigsohn, Hermann. Sitzungsber. Ges. naturforschender Freunde, Berlin.
'l867.

Joint Secretaries. Minutes of the Judicial Commission Meetings held in Rome
(1953) . Internatl. Bull. Bact. Nomen. and Tax. 3 (4) :151. 1953.

Judicial Commission. Opinion 13. Ibid. 4 (3-4) :151-156. 1954.

Kurth, H. Ueber Bacterium zopfii, eine neue Bakterienart. Ber. Deut. Bot.
Ges. 7:97-100. Berlin. 1883.

760 Infernafional Code of Nomenclature

Appendix C. Opinions 13, 14.

Mueller, O. F. Vermium terrestrium et fluviatilum seu animaliuin infusorior-

uni, helminthicorum et testaceorum, non marinorum, succincta historia.

Vol.7. 1773.
Pacini, Filippo. Osservazioni microscopiche e deduzioni patologichc sul

cholerae asiatico. Gazz. med. ital. feder. tosc, Firenze. 1854.
Rasmussen, Anker F. Om dryckning af Microorganismer fra spyt of sunde

mennesker. Copenhagen, 1883. (Dissertation) .
Robin, C. Histoire naturelle des Vegetaux parasites qui croissant sur 1' Homme

et sur les Animaux vivants. p. 345. Paris. 1853.
Sand, G.. and C. O. Jensen. Die Aetiologie der Driise. Deut. Zeitschr. f. Thier-

med. 73:437-464. 1888.
Trevisan, Vittore. Prospetto della Flora Euganea. 1842.

. Nomen. Alg. 1845.

. Prime linee d'introduzione alio studio dei batteri italiani. Rendiconti,

Real Istit. Lombaro dei Scienze e Letter. Ser. II, 72:133-151. 1879.
. Caratteri di alcuni nuovi generi di Batteriacee. Atti della Accad.

Fisio-Med.-Stat. Milano. Ser. IV, 5:92-107. 1885.
. II fungo del cholera asiatico. Questioni risolto. Atti Accad. Fisio-Med.

Stat., Milano. Ser. IV. 5:78-91. 1885.
. Sul micrococco della rabbia e siilla possibilita di riconoscere durante

il periodo incubazione, dall 'esame del sangue della persona morsicata,

se ha contratta I'infezione rabbica. Rendiconti Reale Istit. Lombardo di

Sci. e Lettere. II. 20:88-105. 1887.

I Genera e le Specie della Batteriacee. Milano. 1889. Reprinted in

Internatl. Bull. Bact. Nomen. and Tax. 2 (1) : 13-44. 1952.
Vaucher, Jean Pierre. Histoire des Conlerves deau douce. 1903.
Zopf, W. Die Spaltpilze, 3 Aufl. 1885.

OPINION 14

Names of Bacterial Genera to be Rejected as later
Homonyms of Names of Genera of Protozoa

Becker (1951) listed the generic homonyms in bacteriology and proto-
zoology. He found that ten bacterial generic names proposed (one
of them, Astasia, independently for two different bacterial genera)
are later homonyms of names of protozoan genera. He suggested that
these bacterial generic names should be rejected in bacteriology. The
Editorial Board (1953) proposed that these names be considered by
the Judicial Commission for possible inclusion in the list of names
of bacterial genera to be rejected. The proposal has been considered
by the Judicial Cominission and the following Opinion approved.

Opinion 14.

The following names proposed for bacterial genera are found to be later
homonyms of names applied to genera of protozoa. Rule 24 of the Inter-
national Code of Nomenclature of Bacteria and Viruses states that such later
homonyms are illegitimate in bacteriology. These names are to be placed in
the list of names of bacterial genera to be rejected (nomino generum bac-
teriorum rejicienda).

Appendix C. Opinion 14.

Bacteria and Viruses

161

Rejected Names of Bacterial
Genera

Astasia Meyer 1897
Astasia Pribram 1929
Castellanella Pacheco and

Rodrigues 1930
Charon Holmes 1948
Coccomonas Orla- Jensen 1921
Listerella Pirie 1927
Pahnula Prevot 1938
Pfeifferella Buchanan 1918
Phytomonas Bergey et al. 1923
Rhizomonas Orla-Jensen 1909
Rhodosphaera Buchanan 1918

Names of Protozoan Genera
Having Priority

Astasia Ehrenberg 1830

Castellanella Chalmers

1918
Charon Karsch 1879
Coccomonas Stein 1878
Listerella Jahn 1906
Pahnula Lea 1833
Pfeifferella Labbe 1899
Phytomonas Donovan 1909
Rhizomonas Kent 1880
Rhodosphaera Haeckel 1881

REFERENCES

Becker. Elery R. The legitimacy of certain generic homonyms in bacteriology

and protozoology. Internatl. Bull. Bact. Norn, and Tax. / (3): 102-1 12.

1951.
Bergey. D. H., F. C. Harrison, R. S. Breed, B. W. Hammer, and F. AI. Hun-
toon in Bergey"s Man. Det. Bact., Ed. 1. p. 174. 1923.
Buchanan. R. E. Studies on tlie nomenclature and classification of the Bacter-
ia. \. Subgroups and genera of the Bacteriaceae. J. Bact. 5:27-61. 1918.
. Ibid. IX. Subgroups and genera of the Thiobacteriales. J. Bact. 5:461-

474. 1913.
Chalmers, A. J. The classification of trypanosomes. J. Trop. Med. and Hyg.

27:221-224." London. 1918.
Donovan. C. Kala-azar in Madras, especially with regard to its connection

with the dog and the bug [Conorrhinus) . Lancet, 777:1495-1496. 1909.
Editorial Board. Status of names of bacterial genera that are later homonyms

of names of protozoan genera. Preliminary Statement. File 65, Internatl.

Bull. Bact. Nomen. and Tax. 5 (2, 3) : 109-1 10. 1953.
Ehrenberg, C. G. Beitrage zur Kenntnis der Organisation der Infusorien und

ihrer geographischen Verbreitung, besonders in Sibirien. Akad. Berlin.

Abh. der Math, und Phys. Kl.. pp. 1-88. 1830.
Haeckel. Ernst. Entwurf eines Radiolarien-Systems auf Grund von Studien

der Challenger-Radiolarien. Jenaische Zeitschr. f. Naturwissenschaft. 15:

418-472. 1881-1882.
Holmes. F. O. Order Virales. The filterable viruses, in Bergey's Man. Dei.

Bact.. Ed. 6. pp. 1125-1286. 1948.
Jahn, E. Myxomycetensludien. Ber. Deut. Bot. Ges. 2-/:538-541. 1906.
Joint Secretaries. Minutes of the Judicial Commission Meetings held in Rome

(1953) . Internatl. Bull. Bact. Nomen. and Tax. 5 (4) :15I. 1953.
Judicial Commission. Opinion 14. Ibid. 4 (3-4) :156. 1954.
Karsch. F. Ueber eine neue Einteilung der Tarantuliden (Phryuidae aut.)

Arch. f. Naturgeschichte 7:189-197. 1879.
Kent. W. S. Manual of Infusoria 7:224. 1880.

Labbe. A. Sporozoa, iii Das Tierreich. (Lief. 5) pp. 1-180. 1890.
Lea, I. Contributions to Geology. Philadelphia. 1833.
Meyer. Arthur. Studien iiber die Morphologic und Entwicklungsgeschichte

162 International Code of Nomenclature

Appendix C. Opinions 14, 15.

der Bakterien, Ausgefiihrt an Astasia asteroslxjia A. M. uiul Bacillus liinic-

scens Zopf. Flora. Allg. Bot. Zeitg. 5^:185-248. 1897.
Orla- Jensen, S. Die Hauptlinien des natiirlichen Bakteriensvstems. Zent. f.

Bakl., Par. u. Inf.. Abt. II. 22:305-346. 1909.
. The main lines of the natural bacterial system. J. Bact. 6:26.^-273.

1921.
Pacheco G. and C. Rodrigues. Sobre as caracteristicas bacteriologicas dos

bacillos dvsentericos. Arch. Sao Paulo, Brazil, Inst. Biologico. 5:145-176.

1930.
Pirie, H.J.J. A new disease of veld rodents. "Tiger River Disease." Publ.

South African Inst. Med. Res. 3:163-186. 1927.
Prevot, A. R. Etudes de Systematique Bacterienne. IV. Critique de la con-
ception actuelle cki genre Clostridium. Ann. Inst. Past. 61:12-91. 1938.
Pribram. E. A contribution to the classification of microorganisms. J. Bact.

75:361-394. 1929.
Stein, F .R. Der Organismus der Infusionsthiere. Abt. III. I Hiilfte. Tafel

XXIV. 1878.

OPINION 15

Conservation of the Family Name Enterobacteriaceae,
of the Name of the Type Genus, and Designation of the Type Species

The Editorial Board of the Judicial Commission published a state-
ment relative to a proposal that the family name Enterobacteriaceae
Rahn 1937 be conserved, that the type genus be designated, that the
name of the type genus be conserved, and that the type species be
designated. The Commissioners were requested to \ote by ballot on
the several proposals.

The result of the vote authorizes the Editorial Board acting for
the Judicial Commission to publish the following Opinion. Ihis
Opinion is final unless revoked or modified by action of the Inter-
national Coinmittee on Bacteriological Nomenclature.

Opinion 15.

1. The family name Enterobacteriaceae Rahn 1937 (p. 280) is placed in the
list of conserved family names (nomina conservanda familiorum).

2. The genus Escherichia Castellani and Chalmers 1919 (p. 941) is designated
as the type genus of the family Enterobacteriaceae Castellani and Chalmers
1919.

3. The generic name Escherichia Castellani and Chalmers 1919 (p. 941) is
placed in the list of conserved generic names (nomina conservanda
generum).

4. The type species of the genus Escherichia Castellani and Chalmers 1919
(p. 941) is Escherichia coli (Migula) Castellani and Chalmers 1919 (p. 941),
(basonym Bacillus coli Migula 1895 (p. 27); hyponym Bacterium coli com-
mune Escherich 1885 (p. 518)).

REFERENCES

Editorial Board. Proposal to conserve the family name Enterobacteriaceae
and to designate the tvpe genus. Internatl. Bull. Bact. Nomen. and Taxon.

7 (4): 165.' 1957.

APPENDIX D

Conserved and Rejected Names
of Taxa

APPENDIX D

Conserved and. Rejected Names of Taxa

Nomina Taxorum Conservanda et Rejlcienda

The International Code ot Nomenclature of Bacteria and Viruses
provides for publication of lists of names of taxa of the several ranks
which are to be conser\'ed or rejected.

The conserved and rejected names and specific epithets approved
to January, 1958, are here listed in five tables.

Table I. Conserved and rejected family names.

Table II. Conserved names of genera of bacteria.

Table III. Conserved specific epithets in names of species of bac-
teria.

Table IV. Rejected names of genera of bacteria and viruses.

Table V. Rejected names of subgenera.

CONSERVED AND REJECTED FAMILY NAMES
Nomina familiarum conservanda et rejicienda

Conserved Name Name of Type Genus Rejected Name Opinions

Nomen Conservandum of Conserved Family Nomen Rejiciendum No. and Dat

Enterobacteriaceae Escherichia^ C a
Rahn 1937, p. 280. tellani and Cha

1919, p. &41.

3:74.

195?

3:127.

1953

3:149.

1953

3:153.

1953

3:159.

1953

7:165.

1957

8:73.

1958

»I.B . -International Bulle
Iowa State College Pres

I of Bacteriological Noi
Ames, Iowa. U.S..^.

[ 165

766

International Code of Nomenclature

Appendix D. Conserved Names of Gene

Bacteria and Viruses 167

Appendix D. Conserved Specific Epithets

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Index

a, feminine ending, 32

a, illegitimate as specific epithet, 95

a, becomes ae, 101, 104

a, becomes aa, 101, 104

aa, = a, 101, 104

Abbreviations

of authors' names, 76

of species names, 117
abrotonum, 103
Absence of a Rule, 9
Acarus, 103

Accessory considerations, 9
-aceae, family suffix, 22, 24, 86, 102
Acetobacter, 28, 114, 115

kuetzii^giaims, 110

pasteurianus, 110
Acetomonas, 28
Achrom., abbr., 117
Achromobacter, 102, 114
Achromobacteriaceae, 102
achromogenes, 114
acidophilus, 42, 114
Actino., abbr., 117
Actijiobacillus, 28, 114

mallei, 170
Actinobacter du lait visqueux, 39
Actinomyces, 27, 30, 114, 118, 137

albus, 58

californicus, 37

farciniciis, 166

foersteri, 104

forsteri, 104

nocardii, 166
Actinornycetaceae, 30
aci/5, actinis, 137

Adjectival epithet of name of a man, 109
Adjectival form of a word (for

epithet) , 109
Adjective as specific epithet, 36

formed from name of person, 37

formed from place name, 37
ae = a and a, 101, 104
-ae, feminine genitive ending. 111
ae — ae, 101, 104
aer-, 103
aeris, 103
Aero., abbr., 117
Aerobacillus, 81

poly my xa, 28
Aerobacter, 27

aerogejies, 82

cloacae, 82
Aerobacter aerogenes, 103
aeruginosa, 103
a«, 103

Aesculapius, i, 109
agalactiae sp. ep. cons., 167
Agreement (grammatical) , 33
aguti, 42
af, incorrect transliteration, 137

albicomus, 113
albidoflavus, 114
albochryseus, 42
Albococcus, 81
Albofaciens, 31
albogilvus,36, 113, 114
alboniger, 114
albosporeus, 114
Alcaligenes, 31,95, 118

alcaligenes, 95

faecalis, 95
-rt/<?5, ordinal ending, 24, 86
Algae, departure date, 21, 59
-a/w, meaning of, 24
alpha

as a specific epithet, 95

transliteration of, 137
Alteration

of circumscription of taxon, 81

of diagnostic characters, 75, 81

of status, 81
Alternates

nominations for, 125

recognition of, 125
Alternative spellings

of epithets, 141

of gender and case endings, 144

involving diphthongs ae and oe, 143,
144

of names, 141
Ambiguous names (rejection) , 91, 124
Ambroz, 15

Amendments to Code, 123
American Code of Entomological

Nomenclature, x
American Subcommittee, x
Anaerogenic mutant, 49
Anderson, 91, 151
angium, 137, 139

Anticipation future acceptance, 59
-anus, as suffix to name of a man, 110
-anus, -a, -um, adjectival endings for

names of women. 111
Appendices, definition of, 5
Appendix

A, 135

B, 141

C, 145

D, 163

Application, of names determined by

type, 49
Apus, 90
aquifolius, 113
Aragao,9\, 151
Arbitrary gender, 118
arcesius, ii, 109
Archangium, 137, 139
Argentina, xiv
Artemisia abrotonum, 103
Arthro.,abbr.. 117

[173]

174

International Code of Nomenclature

Index

Arthrobacter, 118

Arthromitus batracliorum, 38

Astasia Ehrenberg 1830, 161

Meyer 1897 nom. rejic, 161, 168
Pribram 1929 nom. rejic, 161, 168

aster ospora, 168

atricapillus, 113

atrojaciens, 114

Aurococcus, 81

aurogenus, 114

Australia, xiv

Author of a scientific name, definition
of, 74

Autograph (eff. publ.) , 61

auxe, 139

auxinophilus, 137, 139

auxo, 137

Available name, definition of, 12

Azoto., abbr., 117

Azotobacter,m, 102, 118
vinelandii, 41

b, illegitimate specific epithet, 95
Babesia Trevisan 1889 nom. rejic, 168

xanthopyrethica Trevisan 1889 Type
sp., 157, 168
Bacillaceae, 53, 54
Bacillariophyta, 153
Bacilleae, 54
Bacillinae, 54
Bacilloideae, 54

Bacillus Cohn 1872, g. cons., ix, 17, 27,
31,53,54,81. 118, 148,166

achrous, 36

aeris, 103

(Aerobacillus) polymyxa, 28

aerofoetidus, 92

albolactis, 36, 115

a/bu5 /ac/w, 35

albus-lactis, 35, 36

alkalescens, 169

a/i/e'? Cheshire and Cheyne 1885, 158,
169, 172

afoef Flugge 1886, 158

aterrimus, 85

(Bacillus) subtilis, 28

campestris, 171

co«, 166

dysenteriae, 167

faecal is alcaligenes, 31

flavoviridis, 36

jluorescens liquefaciens, 44

kaustophilus, 103

ma//e/, 170

megaterium deBary, 101
spelling of, 149

niger, 85

pasteurii, 43

pneumoenteritidis-murium, 41

putrificus coli Flugge 1886, 158, 172

pyocyarieus, 97

type species of Pseudomonas Migula
1894, 150

radicicola, 37

rasmusseni , 168

septicaemiae-anserum-exudalivae, 41

septicaemiae-haemorrhagicae, 35

s^x<u5, 95

iufe/iVw Cohn 1872, ix, 17, 53, 81, 85,
166
aterrimus, 44
niger, 44

subtilis subsp. niger, 44

subtilis subtilis, 44

subtilis var. aterrimus, 85
var. niger, 44, 85
var. subtilis, 85

tertius, 95

thermodiastaticus, 115

tremulus Koch 1877, 158, 172

viscosus, 4 1
BacL, abbr., 117

-bacter, stem (combining form) of,
102, 149

gender of bacterial names ending in,
109, 149
Bacteria, 1 May, 1753, 59
Bacteriaceae, 54

a nomen familiae rejiciendum, 149, 165
bacteriferus, 114
Bacteriological Code, ix

definition of, 3
Bacteriological nomenclature, scope, 13
Bacteriophages, 49

Bacteriopsis Trevisan 1885 nom. re/iV..
158, 168

rasmussenii Trevisan 1885 Type sp.,
158, 168
-bacterium, 119

gender, stem (combining form) of,
102, 149
Bacterium Ehrenberg nom. gen. rejic,
27,31,32,66,92, 118, 137, 149

aeruginosutJi, 97, 102, 166

alcaligenes, 96

articulatum, 170

campestris, 171

chryseum, 37

co/? comjnune, 58, 166

laminariae Billet 1888, 158, 168

mfl//er, 170

inonocytogenes, 166, 170

710 car di, 166

pneumoniae crouposae Zopf 1885 Type
sp., 157, 166

qualis, 95

sonnei, 167

triloculare, 92

zo/7/i/ Kurth 1883 Type sp., 157, 166
Bacteroides, 118

coagulans, 37
-bad roil, stem

(coml)ining form) of, 149
-bactrum, stem (combining form) of, 149
bakterium. 138
Baldwin and Fred, 83
Barkeria, 41
Barton, 108
Bartonella, 74, 108

bacilliformis, 74
Basionym (author citation) , 63
Batteriacee, 24
Becker, 93, 1.56, 161

Index

Bacteria and Viruses

175

Beggiatoa Trevisan 1842 nom. cons., 60.
107, 157, 166

alba (Vaucher) Trevisan 1845 Type
sp., 157, 166

punctata, 166
beigelianum, 43
beigelii, 43

Beijerinck, 65, 70, 103
Belgium, xiv
Beneckea, 107
Bergey, 73, 151, 161
Bergey et al., 73

Bergey's Manual of Determinative Bac-
teriology, 93, 107
Berkeley, 67

beta, transliteration of, 137
Betabacterium, 28, 85
Biblia Latina Vulgata, 116
Bibliographic references; see Citation
Bier, 151
Billet, 24, 159

Billetia Trevisan 1889 nom. rejic, 158,
168

laminariae (Billet) Trevisan 1889
Type sp., 158, 168
Binary combination, 15, 28, 33, 71
Binomen, 34

Binominal combination, 34
Biotype, 45, 49

anaerogenes, 49
Bisby's Dictionarv of the Fungi, 93
Bizio, 73, 91
Blastocaulis, 114
Bollinger, 159
Bollingera Trevisan 1889 nom. rejic, 168

equi (Rivolta) Trevisan 1889 Type
sp., 159, 168
Bordet, 108
Bordetella, 108
Bornet and Flahault, 59
Borrelia, 28, 107

kochii, 38
Bory de St. Vincent, 103, 153
Botanical Code, ix

definition of, 3
Botryomyces ascoformans, 168

equi, 168
Boyd, 154, 1.55

boydii, sp. ep. cons., 155, 167
brachys, 31
Brazil, xiv

Breed, vii, ix, x, xiv, 70, 73, 148
Brevibacterium, 28, 31, 32, 114, 115
British Desmidieae, 59
Brown, x
Bruce. 108
Brucella, 108

mallei, 170

melitensis, 37
Brues, 74

Buchanan, vi, xiii, ix, x, 70, 92, 148, 161
Bulgaria, xiv

Caesarea, 69

Calyjnmatobactenum granulomatis Ara-
gao and Vianna 1913, 91, 151, 169

Cambridge, 115

Canada, xiv

cantabrigensis , 115

Capitalization of specific epithets, 102,

148
Card catalogue of validly published

names of plant genera, 93
Caryo., abbr., 117
Castellanella Chalmers 1918, 161

alcalescens, 169
Castellanella Pacheco and Rodrigues

1930 nom. rejic, 161, 169
Castellani and Chalmers, 31, 32, 96, 155
Category; see Rank
Catenabacterium, 114
caustophilus, 103
cedonulli, 42
cedo-nulli, 42
Cell., abhr., 117

Cennmesia Trevisan 1889 nom. rejic,
158, 169

albida Trevisan 1889 Type sp., 158, 169
cerinus, 138
chaeta, 138
chaete, 138

ChaiiTnan and vice-chairman of Inter-
national Committee, election of, 126
Chairman of International Committee,

duties of, 126
Chairman of Judicial Commission, duties

of, 130
Chalmers, 161
Change of name, 16
Changes

in names as a result of change in
rank, 81

in names as a result of segregation, 81

in names as a result of union, 81
Charon Holmes 1948 nom. rejic, 161, 169
Charon Karsch 1879, 161

evagatus, 169
Cheshire, 159
Chester, 103
Cheyne, 159

chi, transliteration of, 140
Chlamydobacteriales, 140
chlamys, chlamydis, 140
Chloro., abbr., 117
Chlorobacterium Guillebeau 1890 nom.

rejic, 78, 79, 150, 169
Chlorobacterium Lauterborn 1915,

conservation of the generic name,
150

lactis, 169
Cholera of fowls, 35
Chondromyces, 27

crocatus, 67
Chrom., abbr., 117
Chromatium, 65
Chromobacteriaceae, 102
Chromobacterium, 102, 118
Citation of a name published as a syno-
nym, 78

of an infrasubspecific form, 47

of authors, 73

of names, 73
Citing in synonymy, 78
Cladascus mallei, 170

176

International Code of Nomenclature

Index

Clements, 30
Cloaca, 27

Clostridium, 27, 108, 118
botulinum, 36
malenominalutu, 37
oedematis, 92
oedematis-maligni, 36
pasteurianum, 37, 41, 43, 103
perjringejis, 92
sex turn, 95
sp., 95
welchii, 92
Clostrinium, 79
Coccobacillus avicidus, 166
Coccomonas Orla-Jensen 1921 «o;?!. re/zV.,
161, 169
Stein 1878, 161
Code of Bacteriological Nomenclature, x
Code of Ethics, 94
Cohn, 59, 84
Colesiota, 108
Colla, 107
Collaea, 107
comb, nov., 73

"Combination," definition of, 33
Combined generic and specific descrip-
tion, 70
Commission on Nomenclature and

Taxonomy, vi
Commissioners, xi

Committee on Classifioition of the So-
ciety of American Bacteriologists, 60
Commonwealth Mycological Institute,

Kew, 93
Compound epithets, 113
Compound Greek adjective, 37
Compound Latin adjective, 36
Compound names, 100

of taxa, 113
Conn, ix, 148

Conservation of names of genera pro-
posed by Trevisan 1842-1890, 156
generic name Bacillus, 148
Conserved names, 123
of family, 165
of taxa, 165
Conserved specific epithets, 167
cornanguinum, 42
corn-anguinum , 42

Cornilia Trevisan 1889 nom. rejic, 158,
169
alvei (Fliigge) Trevisan 1889 Type

sp., 158, 169
(Eucornilia) alvei Trevisan 1889 Tvpe

sp., 158, 172
(Pleurospora) tremula (Koch) Trevi-
san 1889 Type sp., 158, 172
cornupastoris, 42
corjiu-pastoris, 42
Correct name, 95
definition of, 15
Zoological requirements, 97
coryne, 138

Coiynebacteriaceae, 102
Corynebacterium, 27, 30, 70, 79, 100, 102,
113, 114, 138
bovis, 38
diphtheriae, 48
mallei, 170

coiyiicphorus, 102, 1 1 1

Corynobacterium, 102

Cowan, 154, 155

Cowdria, 107

Cox, 108

Coxiella, 76, 108

Crenothrix, 118

cristalliferus, 114

Criteria employed in classification ,viii

Cultivars, 48

Culture, 56

Curtilius, ii 109

Curtis, 67

Curtius, i, 109

Cuspius, ii, 109

Cyrillic alphabet, 104

Cytophaga krzemieniewskae, 41

Date

of acceptance of an article for publica-
tion, 61

of epithet, 67

of name, 67

of publication of separates, 68
Deazotoriitrauitriazobacterium, 15
De Bary, 101
De CandoUe, 77
Dedication of genera, 107
delta, 95

transliteration of, 138
De Monbreun, 91, 151
Denmark, xiv
De Petschenko, 65, 154
Designation of a type species, 68
Desmidiaceae, 1848, 59
De Toni, 159
Deutsches Reich, xiv
Diacritic signs, 101, 103
Diaeresis ,103
diarrhoea, 139
diarrhoeae, 139

Diagnosis in familiar language, 68
Dicoccia Trevisan 1889 nom. rejic, 158,
169

glossophila Trevisan 1889 Type sp.,
158, 169
Dicrobactrum, 91

Didymohelix Griffith 1853 nom. rejic,
153, 169

ferruginea ,153, 166, 169
Diphasfc variation of the flagella, 51
Diphthongs, 101
Diplococcus, 138

gonorrhoeae, 166
diplus, 138

Disagreeable names, 89
Discomyes, 168

equi, 168

forcinicus, 166
Division of a genus, 26
Division of species, 83
Donker, 81
Donovan, 161

Donovania Anderson, de Monbreun and
Goodpasture 1944 nom. rejic, 107,
169

granulomatis, 91, 151, 169

Incle>

Bacteria and Viruses

177

Dow son, 81

Drepanospira de Petschenko 1911,
validly published, 65, 154
mueUeri, 65, 154, 170
Duly published, 12,61
Dysenteriae sp. ep. cons., 167
Dysenteroides, 31

e. 101, 138

e incorrectly transliterated, 137

-e-, 102

e, 101, 104

e, 101, 104

e, 101, 104

-ea, 24

-eae, 24, 86

Eberthella, 28

Eboracum, 69

echinococcus, 42

Editorial Board, xv, 93, 150, 151, 152.

153, 154, 155, 156, 157, 159, 161
Edwards and Fife, 82
Effective publication, 61, 63
Ehrenberg. 59, 91, 92, 103, 153, 159, 161
Ehringer, 73
Eire, xiv
-ella, 108
emefid., 75
emendai'it, 75
Encapsulatus, 31

inguinalis Bergey et al. 1923, later
synonym of Calymmatobacterium
granulomatis Aragao and Vianna
1913, 151, 169
encephaloides, 137
encephalus, 137
Enderlein, 79, 91, 102
Engler, 59
Enterobacteriaceae , 54

nom. cons., 162, 165
Enterobacteriaceae Subcommittee, 38, 50,

82, 155
entenim, 138

Ephemeral publications, 61
Epithet, 27

definition of, 33

of species, 44

of subspecies which becomes a species,
86
epitheta specifica conservanda, lists of,

133, 167
epitheta specifica rejicienda, lists of, 133
epsilon, transliteration of, 138
Encinia, 28, 107

citrimaculans, 37

lathyri, 38
erysipelatis, 139
Erysipelothrix, 138
Escherichia Castellani and Chal-
mers, 1919, 28, 58, 107, 119, 162

placed in list of nom. cons. gen. 162

type genus of Enterobacteriaceae Cas-
tellani and Chalmers, 1919, 162.
166

type species basonym Bacillus coli
Migula 1895, 162

type species Escherichia coli (Migula)

Castellani and Chalmers, 1919, 162
type species hyponyra Bacterium coli
commune Escherich 1885, 162

Escherichia coli (Migula) Castellani and
Chalmers 1919, type species of
Escherichia Castellani and Chal-
mers 1919, 58, 162, 166

Escherichia group, 50

eta, transliteration of, 138

Etymology

of names of taxa, 69

of new generic names, 1 15

of new specific epithets, 115

eu, 138

eu-, 139

Eubacteriales, 139

Eubacterium, 138

Eucornilia Trevisan 1889 nom. rejic.
158, 172

Eugenis laurina, 78

Eunianlegazzaea Trevisan 1889. iiom.
rejic, 158, 172

Eupacinia Trevisan 1889 nom. rejic,
158, 172

eurectus, 139

eus, 138

Euspirillum Trevisan 1889 nom. rejic,
158, 172

Ewing, 155

ex, 78

Examples, x

Exceptions to the Rules, 123

excl. gen., 75

excl. spec, 75

Falsibacillus, 31
jalsus, 31
Family, 24
"Family Group," 14

termination, 22
Faulty transliteration, correction of, 116
Fischer, 66, 79
Flavobacterium, 28
flavochromogenes, 114
flavogriseus, 114
jlavovirejts, 114
Fleming, 101
Flexner, 155

flexneri, sp. ep. cons., 155
Fliigge, 159

Font used for scientific names, 105
Fore^vord, v
Form, 51
foryna, 45, 51
forma specialis, 14, 45, 51
France, xiv
Frank, 83

Fungi, 31 Dec. 1801,21,59
Fusiformis. 28, 31

Gaffkv, 159

Gaffkya Trevisan 1885, 28, 107, 159
tetragena (Gaffkv) Trevisan 1885
Type sp., 159
Gai, 109'

778

International Code of Nomenclature

Index

Gaillonelle, 103, 153
Gains, 109
i^a Id . ii.(i Id ctis, 137
CaUirtoaxcus, 137

GalUonella Ehrenberg 1838, nom. cons.,
28, 103, 153, 166
ferruginea Ehrenberg, 153, 166, 169
gamma, transliteration of, 137
Gangrenous mastitis of sheep, 35
Gasteromycetes, 58
Gastiaburu, 74
Gender of generic names, 118

names of epithets, 99
Genera united as subgenera under one

generic name, 85
General Consideration No. 1, 3; No. 2,

5; No. 3, 6
General considerations, x
General Principles, x
Generic Group, 14
Generic homonyms in the group

Protista, 148
"Generic/Subgeneric Group," 14
Genitive and adjectival form of same

epithet, 43
Genitive of the name of the genus of
the plant host, 38
singular of a host ,38
singular of a personal name, 38
singular of the name of a locality, 38
Genitives

of names of animal, plural, 38
of names of animal, singular, 38
of names of diseases or lesions, 38
of names of plant taxa in the plural, 38
of other objects, 38
Genotype, 54, 56
Gessard, 97, 150
Glycychylus, 113
Gomont, 59
gonidiaformans, 114
Gonnerman, 78
Goodpasture, 91, 151
Granuloma venereum, 151
Great Britain and Ireland, xiv
Greek, 1 1
Greek stems, 100

long and short, 143
Greek transliteration to Latin, 105
Griffith, 153
griseoflaxnis, 1 1 4
griseoluteus, 114
Group, 14, 45

definition of, 12, 50
Guillebeau, 78. 79, 151
Guillot, 73
gymnocephalus, 42

ha, 137

habrotonum, 103
hae, 137
Haeckel, 161
haema, haematis, 137
haematoides, 137
Haemo., abbr., 117
Haemobartonella canis, 38
Haemophilus, 21, 118, 137

Halnia. 115

Hall,x

Hallier, 67, 92

Ha/o., abbr., 117

halophilus, 137

hals, kalis, 137

Hammer, 73, 161

Hansen, 152

Harrison, 73, 161

Hauduroy, 73

Hauduroy et al., 73

Haupt, 152

Hauser, 73

he, 138

helcogenes, 138

helciis, 138

helianthi, 138

Helikobacterium zopfii, 166

helius, 138

Heller, 32

Hemmi, 110

Hemophilus, 137

Henrillus, 108

Henry, 95

hippopotami, 138

hippiis, 138

Hirn, 59

ho. 139

Hoelling, 31

Holland, xiv

Holman, x

Holmes, 161

Holotype, 58

definition of, 56, 57

Homonyms, 100, 113, 116
of names of genera of Protozoa, 160
in zoology and bacteriology, 93

Hormiscium, 139

hormus, 139

Hucker, 84

Hungary, xiv

Huntoon, 73, 161

Huss, 95

hy, 139

Hyalococcus pneumoniae, 166

Hydro., abbr., 139

Hydrogenomonas, 139

hydrophilus, 139

hyos, 139

Hyphenation of compound specific epi
thets, 36

hys, hyos, 139

138

-:-, as connecting vowel, 100, 113
-ia, 32

-ianus as suffix to name of man, 1 10
-ianus, -a, -um, adjectival endings for

names of women, 1 1 1
ichthyosmius, 138
ichthys, 138
-fdae, 21,22, 24
-iella, 108
-ilia, 108

Illegitimate epithet, definition of, 12
Illegitimate homonyms, xii
Illegitimate name, definition of, 12, 91

Bacteria and Viruses

179

Index

Illegitimate species names, 35

Illegitimate vernacular genitives, 38

-ilium, 108

-illm, 108

-ina, 24

-inae, 22, 24, 86

Inappropriate epithets, 89

Innappropriate names, 89

Incidental mention of a new name. 65

Index Bergeyana, 93

Index Kewensis, 93

Index of Fungi, 93

-inea, 24

-ineae, 24, 86

Infrasubspecific forms, 14, 45, 46

Infrasubspecific names, 14, 46

Infrasubspecific subdivisions, 45

Initial capital, 26

International Association of Micro-
biological Societies, 6, 124

International Association of Micro-
biologists, xi

International Bacteriological Code of
Nomenclature, xv

International Botanical Congress, 1930,
viii

"International Bulletin of Bacteriologi-
cal Nomenclature and Taxonomy,"
XV, 93

International Bureau for Plant Tax-
onomy and Nomenclature, 93

International Code of Nomenclature of
Bacteria and Viruses, xv

International Committee on Bacterio-
logical Nomenclature, 6, 124
functions of, xi. 125
membership of. xi\

International Microbiological Congress
First, vi
Second, ix
Third, xi
Fourth, xiv
Fifth, XV
Sixth, XV

International Rules of Botanical Nomen-
clature, X

International Rules of Zoological No-
menclature, X

International Society for Microbiology,
vi, vii

Interpretation of Rules, 123

Invalid names of genera, 70

-iota, 108

iota, transliteration of, 138

-is, 139

Italy, xiv

Itzigsohn, 159

Jahn, 161

janmaveni, 42

jan-maveni , 42

Jersey, 69

Joint Secretaries, 151, 152, 153, 154, 155,

159, 161
Judicial Commission, xi, xii, 6, 93, 99,
147, 150, 151, 154, 156, 157, 159, 161
functions of, xii, 131

initial membership of, xiii
authorization of, 130

k in Latinized names, 101
kappa, transliteration of, 138
Karsch, 161
Kent, 161
Kisskalt, vi
Kitt, 35, 152
Klebahn, 59
Klebs, 108

Klebsiella Trevisan 1885 nom. cons., 28,
82, 108, 157, 166
granulomatis Bergey et al. 1925 later

synonym of Calymmatobacterium

granulomatis Aragao and Vianna

I9I3, 151, 169
pneumoniae (Schroeter) Trevisan 1887

Type sp., 157, 166
koactli, 42
Koenig, 78
Kolle, 74
Kraus, 74
Kruse 103,
Krzemieniewska. Ill
krzemienieu'skae. 111
Kuckenthal, 93
Kuetzing, 110
Kuntze, 108
Kurth, 159

Kurthia Trevisan 1885 no7n. cons., 28,
107. 108. 157, 166
laminariae , 168
zopfii (Kurth) Trevisan 1885 Type

sp., 157, 166

Labbe, 161
Lacto., abbr., 117
Lactobacillaceae, 89
Lactobacillus, 28, 85, 89, 118

casei, 38

caucasicus, 85

(Thermobacterium) caucasicus, 40, 8
Lactococcus, 114
Lamarckia, 107

lambda, transliteration of, 138
Later homonym, 91, 95
Lathyrus, 83 '
Latin, II

diagnoses of newly described taxa of
plants, 63

genitive of name of a man ,109

stems, 100

terminations, II, 32, 42
Latinization of names of persons, 108

of personal names, 100
Lauda, 74

Lauterborn, 78, 150, 151
Law of Priority, 60, 95
Lea, 161

Lectotype, definition of, 57
Ledingham, vi
Legitimate epithet, 67, 89

definition of, 12

780

International Code of Nomenclature

Index

Legitimate name, 67, 89

definition of, 12

species name, 35

subspecies names, 36
Lehmann and Neumann, 35, 70, 79, 85,

95, 102
lentiputrescens, 36
Lepto., ahhr., 117

Leptospira icterohaemoglobiniiriae , 41
Leptothrix, 60

buccalis Robin 1853, 157. 159, 166, 171

xantopyreticus Trevisan 1883, 158
Leptothrix I, 168
Leptotrichia Trevisan 1879 nom. cons.,

157, 166

buccalis (Robin) Trevisan 1879 Type

sp., 157, 166, 171
(Leptotrichiella) amphibola Trevisan
1889 Type sp., 158, 172
Leptotrichiella Trevisan 1889 nom. rejic,

158, 172
leptotrichoides. 114
Leuco., abbr., 117
Leuconostoc, 138
leiicus, 138
Levine, 155

Link, 95

Linnaea, 108

Liimaei. 109

Linnaeus, v, 31, 77. 103, 108, 109

Linne, v, 108, 109

lipophagus, 114

Listerella Jahn 1906, 161

hepatolytica, 166, 170
Listerella Pirie 1927 nom. rejic, 156,

161, 170
Listeria Necker 1790, 156
Listeria Pirie 1940, nom. cons., 107, 156,
166

monocytogenes (Murray, Webbe, and
Swann) Pirie 1940, type species
of Listeria Pirie 1940, 156, 166
Location of type strain, 68
Loefflerella mallei, 170
Loehnisium, 104
Lohnisium, 104
Lycopersicon, 139
lysodeikticus, 101, 116
lysodicticus, 101, 116

M

Macintosh, 108
Macintoshillus, 32, 108
Magnus, 59
Magrou, 73
Malleomyces, 92

equestris, 67
Mallius, a, 109

Mantegazzaea Trevisan 1889 nom. rejic,
158, 170
cienkowskii Trevisan 1879 Type sp.,

158, 170
(Eumantegazzaea) cienkowskii Trevi-
san 1879 Type sp., 158, 172
Marburg strain, 148
Maria, 111
mariae, 111
Marmor cruciferarum, 38

megatherium, 101, 149
Membership on International Commit-
tee on Bacteriological Nomenclatuie,
124
Meneghini, 91

Merismopedia gonorrhoeae, 166
Mesnil, ix. 148
Metazoa, ix
Methano., abbr., 117
Meyer, 161
M/cr., abbr., 117
Micrococcus, 84, 138

ascoformans, 168

botiyogenus, 168

cholerae-gallinarum Zopf 1885. 158,
166, 170

equestris, 67

gallicidus, 166

gonococcus, 166

gonorrhoeae, 166

lysodeikticus, 101

mastitidis-gangraenosae-ovis, 35

scarlatinosus Trevisan 1879, 158, 170

sphaericus, 41

tetragenus Gaffky 1883, 159
Microfilm made from manuscripts, 61
Micro7nonospora chalcea, 37
tnicrus, 138

Migula. 79. 81,97, 102, 150
Miyagawa. 108, 109
Miyagawae, 109
Miyagawanella, 108
Modification and amendment

of Code, 123

of Rules, 123
Molisch. 70, 110
Monas prodigiosa, 91
moniliformis, 113
Monographie des Oscillaries, 59
Monographic und Iconographie der

Oedogoniaceen, 59
Monotypic new genus, 70
Morphological type, 49
Morphotype. 45. 49
Morphotype capsulatus, 49
mu, transliteration of, 138
Mueller, vi, 59, 60, 160
Miillerina de Petschenko 1910 nom.
rejic, 65, 170

paramecii, 65, 154. 170
Murray, x
Mutant, 45
mut. char., 75
myces, mycetis, 139
-mycetes, 21
-mycetidae, 21
My CO., abbr., 117
Mycobacterium

mallei, 170

tuberculosis avium, 44

tuberculosis var. avium, 44

tuberculosis var. M. avium, 44
My coder ma, 118
Mycoplana, 139
-mycota, 21
-mycotina, 21
Myrtus serratus, 78
myxa, 138

Bacferia and Viruses

181

Al\xo., abbr., 117
Mxxohacter, 95

aureus, 95
Myxococcus,!^, 138
Myxomycetes, 1 May 1753, 59

N

Naked name, 79
Name

based on bacteriophage abnormalit\ ,
92

cited as synonym, 67

conserved so as to exclude its type, 73

definition of, 12, 66

mentioned incidentally, 65

not accepted by the author, 64

not validly published by mention of
subordinated taxa, 67

of species, 33

of species validly published, 71

of subgenus, position of, 40

of subgenus which becomes a genus, 86

of subspecies validly published, 71

of suprageneric taxon changed in
rank, 86

on culture not effective publication, 64

proposed in anticipation of future ac-
ceptance, 65
Names

between suborder and genus of taxo-
nomic groups, 22

of genera and subgenera, definition of,
26

of subgenera, 28

of subspecies, 44

of taxa, 21

of taxa between subclasses and genus,
24

of the Family-Group of categories, 86
National Microbiological Societies, xiv
Naumann, 108
Naumanniella, 108
nc, 137
nch, 137
Necker, 156
necrophorus, 138
necrus, 138
ng, 137

Neisseria Trevisan 1885 nom. cons., 107,
157, 166

gonorrhoeae Trevisan 1885 Type sp.,
157, 166

meningitidis group I Scott, 50

sicca, 36
Neotype culture, 56

definition of, 57
Xevskia, 107
New combinations, 73
New specific (or other) epithets, 112

how written, 113
New York, 115
n igro macu lans, 1 1 4
nigromaculatus, 36
Mtro., abbr., 117
Nitrobacter, 102
Nitrobacteriaceae, 102
Nitroso., abbr., 117

Nocardia Trevisan 1889 nom. cons., 107,
166
alba, 166
caprae, 38
farcinica, 166
maculata, 37
Noguchi, 109
Noguchia, 107
Noguchii, 109
nom. cons., 78
nomen

ambiguum, 92
consenandum. , 78
definition of, 79
dubium, 91

generis conservandum, 79
novum, 78
nudum, 78
Nomenclatural rules, 21; see also Rules
of Nomenclature status of infrasuh-
specific forms, 46
superfluous names, 91
tvpe, 46, 64

definition of, 17
type of taxon of higher rank than a
genus, 53
Nomenclaturally superfluous names, 91
Nomenclature Committee for the Inter-
national Society for Microbiology,
vii
Nomenclature, definition of, 12
nomina ambigua, 91, 124
confusa, 124
conservanda, xii, 15
lists of, 123, 132
conservanda ^amiliarum, lists of, 133
conser-uanda ordinum, lists of ,133
dubia, 92, 124
familiarum conservanda, 165

rejicienda, 165
generica conset-uanda, lists of 132

rejicienda, lists of 133
generum rejicienda, 79, 168, 170
generum virum rejicienda, 168
nomina hybrida, 31, 115
nomina rejicienda xii, 91, 92

lists of, 124
nomina subgenerurn rejicienda, 172
nomina taxorum conservanda et rejici-
enda, 165
Nominations for membership from bio-
logical societies, 124
nom. niid.. 78
"non," 78

Non-declinable Latin noun, 116
Norway, xiv

Nostocaceae Heierocysteae, 1886-88, 59
Nostocaceae Homocysteae, 59
Notes, X

definition of, 5
novaecaesareae, 69
noveboracensis, 69, 115
Novillus. 108
Novu?7i Eboracum, 69
novus, 95
Novy, 108

nu, transliteration of, 138
nx, 137

782

Infernational Code of Nomenclafure

Index

-0-, as connecting vowel, 100, 114

in Greek compounds, 113
o, 101, 104
0, 101, 104

Octopsis Trevisan 1885 7iorn. rejic, 158
170

cholerae-gallinarum Trevisan 1885
Typesp., 158, 170
oe = 6, 0, 101. 104, 138
oedema, 138
oedema maligntim, 36
oedematiens, 138
Oedogoniaceae, 1900, 59
Offensive names, 90
Official Type Culture Collections, xii
-oidae, 24
-oideae, 24, 86
Oldenlandia, 156
omicron, transliteration of, 138
-on, 139

Opinion A, 148; B, 148; C, 148; 1, 149;
2, 149; 3, 149; 4, 149; 5, 149; 6, 150;
7, 151; 8, 152; 9, 153; 10, 154; 11,
154; 12, 156; 13, 156; 14, 160
Opinions, xii, 145

function of, 147

issued by Judicial Commission, 133

request for, xiii
-opsida, 21

"Order/Class Group," 14
Order to genus, 21
Orders, 24

Orders and suborders, names of new, 21
Ordinal adjective, 95
Original spelling, definition of, 99
Orla-Jensen, 85, 162
Orthographic errors, 99

definition of, 101
Orthographic variants, 91, 99, 100, 113,
114, 141

in nomina hybrida, 113
Orthography

of generic names, 29

of names of epithets, 99
-OS, 139

OscilL, zhhr., 117
Oscillaria alba Vaucher 1803 Type sp.,

157
OsciUatoria alba, 166
oum, 140
ozaena, 138
ozaenae, 138

Pacheco, 162

Pacini, 160

Pacinia Trevisan 1885, 159

cholerae-asiaticae Trevisan 1885 Type

sp., 159
(Eiipacinia) putrifica Trevisan 1889

no77i. rejic, 158, 159, 172
(Pseudospira) chlerae-asiaticae Trevi-
san 1885 Typesp., 158, 172
Pack, 104

Pagination of reprints, 68
Palestine, xiv

Palmula Lea 1833, 161

Palmula Prevot 1938 nom. rejic, 161, 170

spermoides, 170
Para., abbr., 117
Parachromatium, 65
Parenthetical citation of author, 76
Park-Williams, 48
Participial adjective

from past participle, 37

from present participle, 37
Past., zhhr., 117
Pasteur, 108, 110

Pasteiirella Trevisan 1887 nom. cons., 28,
103, 108, 119,157

avicida, 166

cholerae gallinarum Trevisan 1887,
Type sp., 35, 157, 166

ellingeri, 73
Pasteuria, 107
pasteurianus, 103, 110
pasteurii, 103
Pastor, 103
Pederson, 28, 85
Pediococcus cerevisiae, 38
Pedioplana

hdckelii, 104

haeckelii, 104
Pe/o., abbr., 117
Permanent secretaries, vii, xi, xiv

election of, 126

duties of. 127
Permanently attached, 53
Perroncitoa Trevisan 1889 no7n. rejic,
158, 170

scarlatinosa (Trevisan) 1889 Type sp.,
158, 170
persicon, 139
Persoon, 59
Petruschky, 31

Pfeifferella Buchanan 1918 nom. rejic,
92, 161, 170

mallei, 170
Pfeifferella Labbe 1899, 161
Pfeiler, 35
Phage type, 49
Phagotype, 45, 49
Phagus, 140
phagus, 140
Phase, 14,45,51
phi. transliteration of, 140
Philip, 76
philus, 139
-phyceae, 21
-phycidae, 21
"Phylum Group," 14
Physiological type, 49
-phyta, 21
-phytina, 21
Phyt07nonas Bergey et al. 1923 nom.

rejic, 161, 171
Phytoninnas Donovan 1909, 161

ca7npestris, 171
pi, transliteration of, 139
Pirie, 156, 162
planus, 139
Plectridixun, 27, 95
Pleurospora Trevisan 1889 no7n. rejic,

158, 172
Plini, 109

Index

Bacteria and Viruses

183

Plinius, 109
Plocamobacteriales, 73
Point of departure for bacterial nomen-
clature, viii
Poland, xiv
Polyangiuni, 60, 95, 118

vitelUnum, 95
Pompei, 109
Pompeius, 109
Prevot, vi, 73, 95, 162
Pribram, vi, 67, 73, 162
Prickett, 103

Principle, No. 1, 9; No. 2. 9; No. 3, 9;
No. 4, 11; No. 5, 12; No. 6, 13; No.
7, 13; No. 8, 15; No. 9, 15; No. 10,
16; No. 11, 17; No. 12, 18
Principles, x, 5, 7
definition of, 5
Priority, 15, 97
Pro., abbr., 117
pro. syn., 78

Propio7iibacteriaceae, 30
Propionibacterium, 30
"Proposals Relative to Emendation and

Publication," xv
Proposed Bacteriological Code of No-
menclature, xiv
Proshigella sonnei, 167
Proteus, 27, 73
Protista, ix

generic homonyms in, 148
generic homonyms not permitted in,
148
Protococcus imetrophus, 91
Protozoa, homonyms of names of genera

of bacteria, 160
Provision for Exceptions to the Rules
and for the Interpretation and
Modification of Rules, 123
Provision No. 1, 123; No. 2, 123; No. 3,

124; No. 4, 124
Provisions, definitions of, 6
pseudes, 140
Pseudobacillus, 31

Pseudomonas Migula 1894 nom. cons.,
27,79.81,97, 102, 118, 140, 166
aeruginosa, 97, 102, 150, 166
barker i, 41
barkeriae, 41
campestris, 171
conjac, 37
cyanogenes, 37
polygoni, 38
pyocyanea, 166
tonel liana, 110
woodsiae, 41
u'oodsii, 41
Pseudospira Trevisan 1889 nom. rejic,

158, 172
psi, transliteration of, 140
Publication

after 1950 of infrasubspecific forms, 46
before 1951 of infrasubspecific forms,

46

of names, 59

Pure suffixes, 108

pus, 138, 139

Putribacillus, 28

qualis, 95

Rahn, x
Ralfs, 59

Ramibacterium, 1 14
Rank of new taxon, 68
Ranks of categories of taxa, 12
Rasmussen, 160

Rasmussenia Trevisan 1889 nom. rejic,
159, 171
anceps, 168

buccalis (Robin) Trevisan 1889 Type
sp., 159, 171
Recommendations, x, 19, 21

definitions of, 5; 2a, 21; 5a, 30; 6a, 40;
6b, 40; 6c, 43; 8a (1) . 48; 8a (2) .
49; 8a (3), 50; 8a (4) , 51; 8a (5) ,
51; 8a (6), 51; 9a, 58; 9b, 58; 12a,
68; 12b. 68; 12c, 68; 12d, 68; 12e,
68; 12f, 69; 13a, 70; 15a, 75; 15b,
75; 16a, 78; 16b, 78; 16c. 78; 16d.
78; 16e, 78; 24a, 93; 27a, 105; 27b,
105; 27c. 105; 27d, 107; 27e, 109;
27f, 111; 27g, 112; 27h, 112; 27i,
113; 27j, 115; 27k, 115; 271, 116;
27m, 116; 27n, 116; 27o, 116; 27p,
117
Rejected names
of families, 165
of genera, 79
of subgenera, 172
of taxa, 165
Rejection and replacement
of names, 89

of names of genera proposed by Trevi-
san 1842-1890, 156
Reprints, 61
Revision des Nostocacees heterocystees,

59
rh, 139

Rh., zhhr., 117
Rhab., abbr., 117
Rhabdomonas fusiformis, 36
Rhizobium, 118

leguminosarmn, 38, 83
phaseoli, 38
Rhizomonas Kent 1880, 161
Rhizomonas Orla-Jensen 1909 nom.

rejic, 161, 171
Rhizopus, 138, 139
rho, transliteration of, 139
Rhodo., abbr., 117
rhodochrous, 139
Rhodococcus, 70

Rhodosphaera Buchanan 1918 nom.
rejic, 70, 161, 171
capsulatus, 171
Rhodosphaera Haeckel 1881, 161
Rhodospirillum, 118
molischianum, 110
rhodum, 139
Ricketts, 108
Picket ts., abbr., 117

184

International Code of Nomenclature

Index

Rickettsia, 76, 107
akari, 103

(Dermacentroxenus) rickettsii, 40
(Rickettsia) proivazekii, 40

Rickettsiella, 108

Ricolesia, 119

Robin, 160

Rodrigues, 162

Ronet, 73

Rosacea, 24

Rosenbach, 81, 84

roseoflavus, 114

Ross, 153, 154

Rough variants, 51

Roumania, xiv

Royal Society, 106

Rubiaceae, 156

rubropertinctus, 114

rudis planusque, 42

Rules, X

definition of, 5

Rules of Bacteriological Nomenclature
with Recommendations, x

Rules of Nomenclature, 19, 21; No. 1
21; No. 2, 21; No. 3, 22; No. 4, 24
No. 5a, 26; 6, 33; 7, 44; 8, 45; 9a, 53
9b, 53; 9c, 54; 10, 59; 11, 61; 12a, 63
12b, 64; 12c, 65; 12d, 67; 12e, 67
12f, 67; 13, 70; 14a, 71; 14b, 71
15a, 73; 15b, 73; 16, 76; 17a, 81;
17b, 81; 18a, 83; 18b, 83; 19, 84:
20, 85; 22a, 86; 22b, 86; 22c, 86;
24, 91; 25, 95; 26, 95; 27, 99; 28. lie

"Rules of Nomenclature, Annotated," x

Saccharo., abbr., 117
Saccharobacilliis pastorianus, 103
St. John-Brooks, vi, vii, ix, xiv
Sale of printed matter, 61
Sahnon, 108
Salmonella, 28, 108, 116, 117

choleraesuis, 38

lexington, 37

london, 37, 116

schottmuelleri, 104

schottmillleri, 104

tennessee, 37
sanctaecatharinae , 42
sanctae-catharinae, 42
Sand, 160
Sapr., abbr., 117
Sarcin., abbr., 117
Sarcma, 27, 60, 118
Sarcinococciis, 70, 115
Saxifraga, 45

aizoon, 45

aizoon subforma surculosa, 45
Schaudinn and Hoffmann, 76
Schizomycetes, 139
Schroeter, 97, 102, 150
Schuetzia, 104, 159
Schulze, 93
Schutzia, 104

poelsii Trevisan 1889 Type sp., 159
Scientific name of an animal, 11
Scientific name of a species, 28
Scientific names, 11
Sclerothrix mallei, 170

Sectio, 14

Section, 14, 27

Seguin, 92

Sellards, 74

Selysius, 107

Separates, 61

Serial letters, as a, b, c not acceptable as

specific epithets, 95
Series, 14, 27
Serological type, 49
Serotype, 14, 45, 49
Serratia, 60, 91, 107

marcescens, 73, 91
Sette, 91
Sextus, 95
Shiga, 108, 109, 155
Shigella, 108, 117, 154

alkalescens, 169

boydii Ewing 1949 validly published,
155, 167

dxsenteriae (Shiga) Castellani and
Chalmers 1919 valid and legiti-
mate, 155, 167

flexneri, 167

flexneri serotype la, 49

sonnei, 167
Shigella, nomenclature of species of

genus, 154
Shigella Center, Chamblee, Georgia, 155
Shigella Commission of the Enterobac-

teriaceae Subcommittee, 154
Shigella group, 50
Sider., ahbr., 117
sigma, transliteration of, 139
Simons, 108
Simonsiella, 108
Simple Greek adjective, 36
Simple Latin adjective, 36
Single specific epithet, 71
Smith. 85
smithae, 111
smithianus. 111
smithii. 111
Sonne, 109

sonnei sp. ep. cons., 109, 167
Spain, xiv
Special forms, 51

species and genera conservanda, viii
species inquirendae, 54
Species name, 34
Species Plantarum, Linnaeus, 1753, viii.

59,60
Species and subspecies names coordinate,

44
Specific epithet, 34

definition of, 33

from name of a man, 109

illegitimate, 95

kinds of, 36

not admissible, 42

not capitalized, 112

recommendations in forming, 40
"Specific name," 34
Specific substantive names, ix
Spelling of Bacillus megaterium deBary,

149
Spelling of synonyms, 78
Sphaer., abbr., 117
sphaericus, 139

Index

Bacteria and Viruses

185

sphaira, 137
Sphinx, 137
sphinx, 137
Spm, abbr., 117
spira, 139
Spirillaceae, 24
Spirillaceen, 24
Spirillacees, 24
Spirillazeen, 24
Spirillum, 24, 27

(Euspirillum) undula (Mueller)

Ehrenberg 1830 Type sp., 158, 172

(Pseudospirillum) amphibolum Trevi-
san 1889 Type sp., 158, 172
spirit us asper, 139
Spirochaeta, 118, 138, 139

pallida, 76
Spironema, 76

pallidum, 76
Sporo., abbr., 117
Si., abbr., 117
Stage, 14, 45, 51
Staphylococcus, 81, 84

ascoformans, 168

aureus, 36, 40

aureus var. e(^u?, 168
State, 14,45,51
Stein, 162
Steinhaus, 95
Stella. Ill
stellae, 1 1 1
Sternberg, 35
Steudel, 78
Sir., abbr., 117
Strain, 14, 45, 48
Strepto., ahbr., 117
Streptobacterium, 28, 85
Streptococcus, 27, 35, 89

acidi-lactici, 35

agalactiae Lehmann and Neumann, 35,
152, 167
710 m. cons., 152

agalactiae contagiosae, 35, 152, 167

cremoris, 38

equi Sand and Jensen 1888, 159

erysipelatis, 103

erysipelatos, 103

group A Lancefield, 50

lactis-acidi, 35, 36

mastitidis, 167

nocardi, 167

of bovine mastitis, 152

pyogenes Rosenbach group A Lance-
field, serotype I Griffith, 50

rubiginosus, 170

xanthopyreticus, 168
Streptomyces, 58

alboflavus, 36

albus, 58

griseus, 48

novaecaesareae, 38, 115

of New Jersey, 115

phaeochromogenes, 37
Streptothrix

alba, 58

farcini, 166

farciui bovis, 166

nocardii, 166
Strong, 74

Subdivisions of species, 45

Subdivisions of subspecies, 45

Subfamilies, 24

subforma. 45

Subgenus, 27

Suborder, 24

Subphyluni, 14

Subsectio, 14

Subsection, 14, 27

Subseries, 14, 27

Subspecies, 45

Substantives

as specific epithets in apposition, 37
in the genitive as specific epithets, 38

"Sufficient nomenclaturally," 12

"Sufficient taxonomically," 12

Suljo., abbr., 117

Superclass, 14

Sweden, xiv

Switzerland, xiv

Symbiotic phage, 49

Synmigium sessile, 36

synchyseus, 137

synchysis, 137

Synopsis Methodica Fungorum, 59

Syntype, definition of, 57

Systema Naturae of Linnaeus, 60

tau, transliteration of, 139
Tautonomy, 55
Tautonym, 95, 96
Taxa, naming of, 21
Taxon, 27

definition of, 12
Taxonomic group, 12

definition of, 15, 21
Taxonomic Subcommittees, functions of,

128
Terms which denote the categories of

taxa, 12
Ternary combination, 15, 44, 71
tetra, 139
tetragenus, 139
th. 139

Thaxter, 78, 97
Thermo., abbr., 117
Thermobacterium, 28, 85
thermofuscus, 114
thermophilus, 36
theta, transliteration of, 138
T/2to., abbr., 117
ThiobaciUus, 118

thioparus, 70
Thioploca, 138
Thiosarcina. 118
Thiothrix, 27
thium, 138
thrix, 138
Tonelli, 110
Transfer of species, 83
Transliteration

of Greek words for use in Nomencla-
ture, 135, 137

of words from languages not using
Latin alphabet, 116
Treponema, 27, 76, 118

calligyrum, 37

pallidum, 76

786

International Code of Nomenclature

Index

Trevisan, 24, 67, 103, 156, 157, 159, 160

Tribes, 24

Trivial name, 34

Trutta trutta, 55

Type, 49

Type culture, 49, 58

Type culture collections, viii

Type culture maintenance, 58

Type genus of a contained family, 21

"Type" material, 58

Type selection by monotypy, 54

by original designation, 54
Type species, viii, 49, 54

definition of, 56

of genus, 53
Type or standard culture, ix
Types are property of science, 58
Typescripts, 61
Typographic errors, 99

definition of, 101
Tyzzer. 74

U

u, 139

u — ue, 101, 104

ue = ii, 101, 104

Uhlenhuth, 74

-urn, 139

Umlaut, 104

Uninomial, 27

Unintentional orthographic error, 99

Unintentional typographic error, 99

Union of Soviet Socialist Republics, xiv

Union of species as subspecies, 85

Union of taxa, 84

United States of America, xiv

upsilon, transliteration of, 139

Urbain, 73

Uredinales, 59

t/ro .,abbr., 117

Urobacillus pasteurii, 103

urubitinga, 42

Uruguay, xiv

-us, 139

U stilagmales , 59

Valid name, definition of, 12

Valid publication, 59

"Validly published," 12, 63

VanLaer, 103

Variant, 14, 45, 48

Variety, 45

Variety and subspecies, 44

Vaucher, 160

Veillon, 108

Veillonella, 108

Viainia,91, 151
Vibrio, 27, 118

berolinensis, 37

cholerae, 158, 172

comma, 37

undula, 158, 172
Vice-chairman of International Commit-
tee, duties of, 126
Vuillemin, 59, 60, 76

W

w in Latinized names, 101
Weinberg, 92
Weldin, 155
Wesenbergus, 32
Winogradsky, 103

Winogradskya Trevisan 1889 riom. rejic,
159, 171
ramigera (Itzigsohn) Trevisan 1889
Type sp., 159, 171
Winslow and Rogers, 81
Wolbach, 108
Wolbachia, 108
Wolhyn, 108
Wolhynia, 108
Wrong connecting vowel, 99

Xanthomonas, 81

antirrhini, 38

campestris, 41, 171

hemmiana, 110

malvacearum, 38

trarislucens i. sp. phlei-pratensis, 51
XI, transliteration of, 138

y, in Latinized names, 101
Yamasaki, vi

Zaogalactina imetrofa, 91

zea, 138

zeae, 138

zeta, transliteration of, 138

Zinssera, 107

Zoogloea, 118

beigeliana, 37

pulmonis equi, 159, 168

ramigera, 159, 171
Zoological Code, definition of, 3
Zoological Record, 62, 93
Zopf, 60, 70, 160
Zopfius zopfii, 166
zoum, 138
Zy mo., abbr., 117

pa

era