IN T ERNA TIONA L B U L L E TIN O F B AC T ERIOLOGICAL
NOMENCLATURE AND TAXONOMY
Vo1.13, N o . 2
A p r i l 15, 1963
pp. 95-110
AN OUTLINE O F NOMENCLATURE F O q
T H E FAMILY ENTEROBACTERIACEAE
W.H. Ewing
Communicable D i s e a s e C e n t e r ,
P u b l i c Health S e r v i c e , U . S. D e p a r t m e n t
of Health, Education, a n d W e l f a r e ,
Atlanta, G e o r g i a
SUMMARY. A n o m e n c l a t u r a l s y s t e m f o r t h e f a m ily Enterobacteriaceae, including four tribes
and ten genera, is presented in outline f o r m
as follows:
T r i b e I. E s c h e r i c h i e a e B e r g e y , B r e e d and M u r r a y . G e n e r a : E s c h e r i c h i a C a s t e l l a n i a n d C h a l m e r s (Type s ~ . g: . k i
(Migula) C a s t e l l a n i a n d C h a l m e r s ) ; Shigella C a s t e l l a n i a n d
C h a l m e r s (Type s p . : 5. d y s e n t e r i a e (Shiga) C a s t e l l a n i a n d
Chalme r s).
T r i b e 11. S a l m o n e l l e a e B e r g e y , B r e e d a n d M u r r a y .
G e n e r a : S a l m o n e l l a L i g n i k r e s (Type s p . : 5. c h o l e r a e - s u i s
(Smith) We ldin); A r i z o n a Kauffmann a n d E d w a r d s (Type s p . :
A . a r i z o n a e Kauffmann a n d E d w a r d s ) ; C i t r o b a c t e r W e r k m a n
a n d G i l l e n (Type s p . : ,C. f r e u n d i i ( B r a a k ) W e r k m a n a n d
G i l Len).
T r i b e 111. K l e b s i e l l e a e T r e v i s a n . G e n e r a : K l e b s i e l l a
T r e v i s a n (Type s p . : 5 . pneumoniae ( S c h r o e t e r ) T r e v i s a n ) ;
A e r o b a c t e r B e i j e r i n c k (Type s p . : A. a e r o g e n e s ( K r u s e )
B e i j e r i n c k ; S e r r a t i a B i z i o (Type sp. : 2. m a r c e s c e n s B i z i o ) .
T r i b e IV. P r o t e e a e C a s t e l l a n i a n d C h a l m e r s . G e n e r a :
P r o t e u s H a u s e r (Type s p . : P r o t e u s v u l g a r i s H a u s e r ) ; P r o v i d e n c i a Kauffmann a n d E d w a r d s (Type s p . : ,P. a l c a l i f a c i e n s
(De S a l l e s G o m e s ) Ewing).
Certain proposals regarding the system a r e
made for consideration by the Judicial Comm i s s i o n of t h e I n t e r n a t i o n a l C o m m i t t e e o n
B a c t e r i o l o g i c a l N o m e n c l a t u r e , a s w e l l a s of
a l l o t h e r s who may be i n t e r e s t e d .
The system is proposed for use by investigators in
R e c e i v e d f o r publication J u n e 14, 1962.
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INTERNATIONAL BULLETIN
a l l f i e l d s of b a c t e r i o l o g y
m e m b e r s of t h i s f a m i l y .
when
dealing
with
The p u r p o s e of t h i s p a p e r i s to p r e s e n t i n outline a s y s t e m of n o m e n c l a t u r e f o r the f a m i l y E n t e r o b a c t e r i a c e a e . The
outline follows the taxonomic s y s t e m previously r e p o r t e d by
Ewing a n d E d w a r d s (I.B.2 lJ:l, 1960) with r e s p e c t to the
v a r i o u s levels of classification.
T h i s taxonomic s y s t e m
found r a t h e r wide acceptance and since it w a s published i n
1960, many have suggested t h a t the author t r a n s l a t e it into
a s y s t e m of f o r m a l n o m e n c l a t u r e that could b e u s e d by the
a u t h o r s of textbooks, j o u r n a l e d i t o r s , and by anyone e l s e
The a u t h o r h a s
who might have need of s u c h a s y s t e m .
a c q u i e s c e d t o t h e s e suggestions and the outline of nomenc l a t u r e to b e p r e s e n t e d i s the r e s u l t ,
I t i s not o u r p u r p o s e to a t t e m p t t o r e v i e w the voluminous
l i t e r a t u r e on the s u b j e c t s of n o m e n c l a t u r e and of taxonomy
within the f a m i l y E n t e r o b a c t e r i a c e a e . Instead, only t h o s e
works t h a t a r e thought to be p e r t i n e n t to the p r e s e n t d i s c u s sion, o r which a r e c o n s i d e r e d i m p o r t a n t s o u r c e s of i n f o r m a tion, a r e quoted. R e f e r e n c e s t o p a p e r s of o r d i n a r y length
a r e i n s e r t e d i n the text, while r e f e r e n c e s to longer p a p e r s ,
monographs, a n d books, a s well a s to s e v e r a l publications
not cited i n t h e text, a r e l i s t e d a t the end. Those i n t e r e s t e d
i n r e a d i n g f u r t h e r on t h e s e s u b j e c t s should consult a f i l e of
the International Bulletin of B a c t e r i o l o g i c a l Nomenclature
and Taxonomy, as well.
The outline i s p r e s e n t e d i n its s i m p l e s t f o r m and this is
followed by a s e r i e s of annotations, i n which p e r t i n e n t f a c t s
a r e mentioned, and i n which attention i s d i r e c t e d to points
that should b e a c t e d upon by t h e J u d i c i a l C o m m i s s i o n of t h e
Nomenclature C o m m i t t e e of the International Association of
Microbiological Societies. R e f e r e n c e s a r e omitted f r o m the
outline but m a y b e found i n B e r g e y ' s Manual, . 6th a n d 7th
editions. F i n a l l y , t h e r e i s a brief discussion.
I . B . = International Bulletin of B a c t e r i o l o g i c a l Nomenclat u r e a n d Taxonomy, Iowa State University P r e s s , A m e s ,
Iowa, U.S.A.
Code = International Code of Nomenclature of B a c t e r i a and
V i r u s e s . 1958, Iowa State University P r e s s , Ames, Iowa,
U.S.A.
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The N o m e n c l a t u r e i n Outline
Annot.
F a m i l y X ENTEROBACTERIACEAE Rahn
1
T r i b e I ESCHERICHIEAE B e r g e y , B r e e d , a n d M u r r a y
Genus I E s c h e r i c h i a C a s t e l l a n i a n d C h a l m e r s
1 . E s c h e r i c h i a &i (Migula) C a s t e l l a n i a n d
Cha l m e r s
Genus I1 Shigella C a s t e l l a n i a n d C h a l m e r s
1 . Shigella d y s e n t e r i a e (Shiga) C a s t e l l a n i a n d
Chalmers
2 . Shigella f l e x n e r i C a s t e l l a n i a n d C h a l m e r s
3. Shigella boydii Ewing
' 4. Shigella sonnei ( L e v i n e ) Weldin
2
2
3
4
T r i b e 11 SALMONELLEAE B e r g e y , B r e e d , a n d M u r r a y
Genus I Salmonella Lignikres
5
1 . S a l m o n e l l a c h o l e r a e - s u i s (Smith) Weldin
6
2 . S a l m o n e l l a typhi ( S c h r o e t e r ) W a r r e n a n d Scott
7
3 . Salmonella enteriditis (Gaertner) Castellani
and Chalmers
8
G e n u s I1 A r i z o n a Kauffmann a n d E d w a r d s
9
1. A r i z o n a a r i z o n a e Kauffmann a n d E d w a r d s
Genus I11 C i t r o b a c t e r W e r k m a n a n d G i l l e n
1. C i t r o b a c t e r f r e u n d i i ( B r a a k ) W e r k m a n a n d
10
Gillen
T r i b e I11 K L E B S I E L L E A E T r e v i s a n
Genus I Klebsiella T r e v i s a n
1. K l e b s i e l l a pneumoniae ( S c h r o e t e r ) T r e v i s a n
2 . K l e b s i e l l a o z a e n a e (Abel) B e r g e y , B r e e d ,
and M u r r a y
3 . Klebsiella rhinoschleromatis T r e v i s a n
G e n u s I1 A e r o b a c t e r B e i j e r i n c k
1. Aerobacter aerogenes (Kruse) Beijerinck
l a . A e r o b a c t e r a e r o g e n e s s u b s p e c i e s hafniae
( M o e l l e r ) Ewing c o m b . nov.
2. A e r o b a c t e r c l o a c a e ( J o r d a n ) B e r g e y , B r e e d ,
and M u r r a y
3 . Ae r o b a c t e r l i q u e f a c i e n s (B e i j e r i n c k ) G r i m e s
and Hennerty
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11
11
12
13
14
15
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Genus I11 S e r r a t i a B i z i o
1. S e r r a t i a m a r c e s c e n s Bizio
la. Serratia marcescens subspecies kiliensis
( L e h m a n n a n d Neumann) Ewing, Davis
and Reavis
T r i b e IV P R O T E E A E C a s t e l l a n i a n d C h a l m e r s
Genus I P r o t e u s H a u s e r
1. Proteus vulgaris Hauser
2. P r o t e u s mirabilis Hauser
3 . P r o t e u s m o r g a n i i (Winslow e_t +,)R a u s s
4 . P r o t e u s r e t t g e r i (Hadley c t 5 1 . ) Rustigian
and S t u a r t
G e n u s I1 P r o v i d e n c i a Kauffmann a n d E d w a r d s
1 . P r o v i d e n c i a a l c a l i f a c i e n s (De S a l l e s C o m e s )
Ewing c o m b . nov.
2. P r o v i d e n c i a s t u a r t i i (Buttiaux e_t
Ewing
c o m b . nov.
16
17
e.)
N.B.
The f i r s t s p e c i e s l i s t e d i n e a c h genus i s the type
species.
Annotations
1. The f a m i l y n a m e E n t e r o b a c t e r i a c e a e Rahn w a s cons e r v e d ( I . B . 8:73, 1958; Code, 165).
2. The n a m e of T r i b e I, E s c h e r i c h i e a e B e r g e y , B r e e d ,
a n d M u r r a y , i s the logical n a m e , s i n c e the genus E s c h e r i c h i a
w a s d e s i g n a t e d t h e type g e n u s of the f a m i l y . ( I . B . 8:73, 1958;
Code, 1 6 6 ; R u l e s 3 a n d 4 , Code, 2 2 , 24).
3. E s c h e r i c h i a &i
(Migula) C a s t e l l a n i a n d C h a l m e r s
w a s d e s i g n a t e d t h e type s p e c i e s of t h e genus ( I . B . 8:73, 1958;
in the species name
Code, 166). T h e s p e c i f i c e p i t h e t &i
(Migula) C a s t e l l a n i and C h a l m e r s should
E s c h e r i c h i a &i
b e d e s i g n a t e d a s a c o n s e r v e d s p e c i f i c epithet (epitheton
s p e c i f i c u m c o n s e r v a n d u m ) f o r the s p e c i e s n a m e d B a c t e r i u m
c o l i c o m m u n e E s c h e r i c h 1885.
It should b e c o n s e r v e d
a g a i n s t t h e s p e c i f i c e p i t h e t e s c h e r i c h i i of t h e s e n i o r (object i v e ) synonym B a c i l l u s e s c h e r i c h i i T r e v i s a n 1889.
An Opinion i s r e q u e s t e d f r o m t h e J u d i c i a l C o m m i s s i o n
of t h e I n t e r n a t i o n a l C o m m i t t e e on B a c t e r i o l o g i c a l Nomenclature.
-
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4 . The n o m e n c l a t u r e (and taxonomy) of genus 11, T r i b e I,
the genus Shigella, h a s b e e n a c t e d upon, a c c e p t e d , a n d cons e r v e d ( I . B . I:147, 1951; 3:82, 1953; 3:150, 1953; &:148,
1954; 8:195, 1958; s : 8 5 , 1960; Code, 167).
5 . Investigation h a s shown that although L i g n i k r e s p r o p o s e d the g e n e r i c n a m e S a l m o n e l l a , he gave no s p e c i f i c
n a m e t o t h e "type le m i c r o b e du hog c h o l e r a de S a l m o n . "
The f i r s t u s e of S a l m o n e l l a i n a l e g i t i m a t e combination w a s
by Buchanan ( J . B a c t e r i o l . 2:53, 1918) who l i s t e d S a l m o n e l l a
a s a subgenus of B a c t e r i u m and a p p a r e n t l y m a d e the f i r s t
typification: " T h e type s p e c i e s i s B a c t e r i u m (Salmonella)
c h o l e r a e suis." H o w e v e r , L i g n i k r e s ' a u t h o r ship of the n a m e
i s not questioned.
6. The c i t a t i o n S a l m o n e l l a c h o l e r a e - s u i s (Smith) Weldin
a p p a r e n t l y i s c o r r e c t . Weldin l i s t e d the c o r r e c t t y p e s p e c i e s .
S a l m o n e l l a c h o l e r a e - s u i s (Smith) Weldin w a s r e c o g n i z e d a s type s p e c i e s of t h e genus i n 1936 (Salmonella Subc o m m i t t e e R e p o r t , J . Hyg. 34:333, 1934; P r o c . 2nd I n t e r n a t .
M i c r o b i o l . C o n g r . , London, 1937, 545).
7 . The s u b j e c t of the s p e c i f i c epithet typhosa v s . the
s p e c i f i c epithet typhi w a s r e v i e w e d by P r o f e s s o r R . S . B r e e d
( I . B . 5-:165, 1955), by the E d i t o r i a l B o a r d ( I . B . 8:155, 1958),
and i s u n d e r c o n s i d e r a t i o n by the J u d i c i a l C o m m i s s i o n . The
taxonomic s u b c o m m i t t e e on the E n t e r o b a c t e r i a c e a e i s on
r e c o r d a s being i n f a v o r of the epithet typhi (I.B. S:173, 1958).
The s p e c i f i c e p i t h e t i n the s p e c i e s S a l m o n e l l a typhi should
be conserved.
8a. If the genus S a l m o n e l l a i s l i m i t e d to t h r e e s p e c i e s ,
a s i n the outline, then a c c o r d i n g to the r u l e s , the n a m e of
the t h i r d s p e c i e s m u s t b e t h a t of t h e o l d e s t validly p u b l i s h e d
s p e c i e s now included i n the genus o t h e r t h a n 2. c h o l e r a e s u i s a n d 5. typhi. The w r i t e r found none e a r l i e r t h a n the
s p e c i e s S a l m o n e l l a e n t e r i d i t i s a s u s e d by G a e r t n e r , 1888.
T h e r e f o r e , the n a m e of the t h i r d s p e c i e s should b e S a l m o n ella enteriditis (Gaertner) Castellani and Chalmer s.
The
e p i t h e t s kauffmannii a n d e n t e r i c a a s u s e d by B o r m a n , S t u a r t ,
a n d W h e e l e r (J. B a c t e r i o l . 2 : 3 5 1 , 1944), a n d Kauffmann
a n d E d w a r d s ( I . B . &:2, l 9 5 2 ) , r e s p e c t i v e l y , a r e l a t e r o b j e c tive synonyms of 5. e n t e r i d i t i s and a r e i l l e g i t i m a t e .
-
3 S e e I . B . Q:31, 1963, Opinion 1 8 .
The s p e c i f i c e p i t h e t
typhi h a s b e e n c o n s e r v e d o v e r the s p e c i f i c epithet typhosa
i n the s p e c i e s n a m e S a l m o n e l l a typhi (Schroe t e r ) W a r r e n
a n d Scott 1930. Ed.
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8b. The concept of limitation of the number of s p e c i e s
i n the genus Salmonella to t h r e e apparently originated with
B o r m a n , Stuart, and Wheeler (loc. c i t . ) .
At least, t h e s e
investigators f i r s t put i t into print, a s f a r a s the w r i t e r h a s
been able to d e t e r m i n e . B o r m a n e t 51. employed the designation kauffmannii f o r t h e t h i r d species of the genus Salmone l l a , However, the s p e c i e s was inadequately c h a r a c t e r i z e d ,
i n the opinion of the w r i t e r , since the c h a r a c t e r i z a t i o n was
based upon antigenic s t r u c t u r e alone. If this c h a r a c t e r i z a tion w e r e used alone many salmonellae could not be readily
classified i n the genus.
Kauffmann and Edwards (loc. c i t . ) a l s o subscribed to
the t h r e e species concept within the genus Salmonella, but
employed the n e u t r a l designation Salmonella e n t e r i c a a s the
name f o r the t h i r d s p e c i e s , which was defined by m e a n s of
b i o c he m i c a 1 c h a r a c t e r i st i c s
8c. The idea of limiting the number of species in the
genus Salmonella to t h r e e is a compromise, which a p p e a r s
to be the only reasonable solution to a difficult problem.
Alternatively, the number of s p e c i e s in the genus could be
limited to one, i n which c a s e the name Salmonella e n t e r i c a
might be conserved f o r u s e i n connection with i t . If this
single species concept w e r e adopted, the names of the v a r i ous s e r o t y p e s could be written Salmonella enterica serotype
typhi o r Salmonella e n t e r i c a (typhi), o r Salmonella e n t e r i c a
(dublin) e t c . However, tradition h a s demanded that the
typhoid bacillus be recognized a s a distinct species and i f
Salmonella typhi i s s o recognized, the type species of the
genus, S. cholerae-suis, a l s o should be given species r a n k .
8d. Other p r o p o s a l s c a l l f o r recognition of the " c l a s s i c a l
species" of Salmonella, These a r e the salmonellae that w e r e
given specific names i n the e a r l i e r d a y s of e n t e r i c b a c t e r i ology and these names were legitimately published. The
principal a r g u m e n t s used i n favor of t h i s proposal a r e that
t h e s e specific n a m e s a r e employed by w o r k e r s in the fields
of b a c t e r i a l physiology, genetics, e t c . , that the antigenic
schema i s not u s e d by physiologists and geneticists, and
that i t i s difficult to integrate the classifications u s e d by the
different groups of investigators. The author i s not i n a g r e e ment with the proposal o r the a r g u m e n t s employed by i t s
advocates. In t h e f i r s t place i t would be impossible to obtain a g r e e m e n t among investigators in the varioils fields and
in v a r i o u s countries on just which should be included i n a
.
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l i s t of " c l a s s i c a l s p e c i e s . ' ' F o r e x a m p l e , the following a r e
l i s t e d i n the 6th edition of B e r g e y ' s Manual (1948):
1. S a l m o n e l l a p a r atyphi (Kay s e r ) C a s t e l l a n i a n d
Chalmers
2 . S a l m o n e l l a S c h o t t m u l l e r i (Winslow g t a l . ) B e r g e y c t 51.
3 . Salmonella typhimurium (Loeffler) Castellani and
Chalme r s
4. Salmonella abortivoequina (Good a n d C o r b e t t )
Bergey gt
5. S a l m o n e l l a a b o r t u s o v i s ( L o v e l l ) Schuetze g t ~ 1 .
6. Salmone lla a b o r t u s b o v i s Kauffmann
7 . S a l m o n e l l a h i r s c h f e l d i i We ldin
8. S a l m o n e l l a c h o l e r a e s u i s (Smith) Weldin
9. Salmonella t y p h i s u i s G l a e s s e r (Schuetze et 5L.)
10. Salmonella m o r b i f i c a n s (Migu l a ) Haupt
11. S a l m o n e l l a typhosa ( Z o p f ) White
12. S a l m o n e l l a e n t e r i d i t i s ( G a e r t n e r ) C a s t e l l a n i a n d
Chalme r s
13. S a l m o n e l l a g a l l i n a r u m (Klein) B e r g e y e_t ~ 1 .
1 4 . S a l m o n e l l a p u l l o r u m ( R e t t g e r ) B e r g e y e_t EL.
1 5 . S a l m o n e l l a a n a t i s ( R e t t g e r a n d Scoville) B e r g e y g t gl.
al.
However, i n the 7th edition (1957), f i v e v a l i d l y p u b l i s h e d
" c l a s s i c a l s p e c i e s " (No. 6, 9, 10, 1 4 and 1 5 above) w e r e not
s o l i s t e d . Also, i t i s known, through y e a r s of c o r r e s p o n dence on the s u b j e c t , that w o r k e r s i n s o m e c o u n t r i e s would
include c e r t a i n o t h e r s a l m o n e l l a e i n s u c h a l i s t a n d would
exclude s o m e of t h o s e l i s t e d above.
Further, many "class i c a l s p e c i e s " cannot b e d i s t i n g u i s h e d f r o m e a c h o t h e r o r
f r o m o t h e r s a l m o n e l l a e by a n y known m e a n s e x c e p t a n t i g e n i c
analysis.
The s t a t e m e n t t h a t b a c t e r i a l p h y s i o l o g i s t s a n d
g e n e t i c i s t s do not u s e the a n t i g e n i c s c h e m a i s i n a c c u r a t e ,
s i n c e m a n y outstanding i n v e s t i g a t o r s i n t h e s e f i e l d s do m a k e
u s e of the a n t i g e n i c s c h e m a . In f a c t , s o m e of t h e m o s t
notable c o n t r i b u t i o n s of the l a s t f e w y e a r s a r e dependent
upon t h e i r u s e of t h e s c h e m a .
8e. Another concept t h a t should b e m e n t i o n e d i s that
a d v a n c e d by P r o f e s s o r Kauffmann, who h a s r e g a r d e d e a c h
Salmonella s e r o t y p e a s a s e p a r a t e s p e c i e s (I.B. 2:1, 7, 1959;
11:5, 1961). T h i s view i s not s h a r e d by m a n y o t h e r s who
r e g a r d t h e n a m e s given t o Salmonella s e r o t y p e s a s i n f r a -
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s u b s p e c i f i c d e s i g n a t i o n s that a r e equivalent to, a n d which
a r e u s e d i n lieu of, the antigenic f o r m u l a s .
(e.g., Internl.
S u b c o m m . E n t e r o b a c t . , I . B . S:178, 1958; L e M i n o r , Ann.
I n s t . P a s t e u r %:207, 1958; Cowan, I . B . 2:131, 1959; T a y l o r ,
I . B . 2:159, 1959). N e v e r t h e l e s s , P r o f e s s o r Kauffmann h a s
e v e r y r i g h t t o h i s v i e w s on t h i s s u b j e c t a n d s i n c e h i s s p e c i e s
a r e d e r i v e d f r o m the a n t i g e n i c s c h e m a t h e r e i s no r e a l
conflict between P r o f e s s o r Kauffmann's v i e w s a n d t h o s e
of m a n y o t h e r s except i n t h e a r e a of the constitution of a
s p e c i e s . However, we a r e of t h e opinion that a n i n t e r p r e t a tion o t h e r than t h a t of P r o f e s s o r Kauffmann c a n b e m a d e of
the s t a t e m e n t of P. B r u c e White ( P r o c . 2 n d I n t e r n . C o n g r .
M i c r o b i o l . 1937, 548) r e g a r d i n g S a l m o n e l l a t y p e s a n d t h e i r
being e l e m e n t a r y s p e c i e s i n the s e n s e of d e v r i e s . F r o m
what t h e a u t h o r h a s r e a d i t s e e m s c l e a r t h a t d e V r i e s cons i d e r e d v a r i e t i e s p r o d u k e d by m u t a t i o n a s e l e m e n t a r y s p e c i e s ; he a p p a r e n t l y c a l l e d s u c h m u t a n t s " e l e m e n t a r y s p e c i e s . " Hence, e l e m e n t a r y s p e c i e s m a y b e c o n s i d e r e d the
e l e m e n t s of a s p e c i e s o r t h e e l e m e n t s that m a k e u p a s p e c i e s ,
r a t h e r than a s s e p a r a t e species.
8f. T h u s , we m a y have one s p e c i e s of S a l m o n e l l a o r
m o r e than 800 s p e c i e s , o r we m a y have a dozen to 20 s p e c i e s of S a l m o n e l l a o r t h r e e s p e c i e s . Anything t h a t i s done
with the n o m e n c l a t u r e of t h e g e n u s Salmonella m u s t b e a
c o m p r o m i s e a n d t h e a u t h o r i s of the opinion t h a t t h e t h r e e
s p e c i e s concept a s p r e s e n t e d i n t h e outline i s the b e t t e r
c h o i c e . S e v e r a l a u t h o r i t i e s i n t h e field a r e i n a g r e e m e n t
with t h i s point of view.
F o r example, D r . P . R . Edwards
a g r e e s with i t a n d h a s s t a t e d ( p e r s o n a l c o m m u n i c a t i o n 1962)
t h a t t h e p r o p o s a l m a d e with P r o f e s s o r Kauffmann i n 1952
(Kauffmann and E d w a r d s , I . B . 2:2,1952) s t i l l i s h i s choice
a s the m o s t l o g i c a l solution to t h i s p r o b l e m .
8g. I t should b e e m p h a s i z e d that i f t h i s p r o p o s a l i s
adopted, attention should b e given to c o r r e c t u s a g e . E x c e p t
w h e r e u s e d i n a g e n e r a l way, a s i n the outline, the n a m e
Salmonella e n t e r i d i t i s should not b e u s e d a l o n e s i n c e i t r e p r e s e n t s 800 and m o r e s e r o t y p e s a n d hence by itself would
b e m e a n i n g l e s s . I n s t e a d , the n a m e of a s e r o t y p e ( o t h e r than
S . typhi a n d 5. c h o l e r a e - s u i s ) should b e w r i t t e n S a l m o n e l l a
e n t e r i t i d i s s e r o t y p e e n t e r i d i t i s o r 5. e n t e r i d i t i s ( e n t e r i d i t i s ) ,
S a l m o n e l l a e n t e r i d i t i s t h o m p s o n o r S. e n t e r i d i t i s (thompson).
I t m a t t e r s not whether the i n f r a s u b s p e c i f i c s e r o t y p e d e s i g nation i s i t a l i c i z e d , s i n c e t h e u s e of i t a l i c s o r o t h e r p r i n t i n g
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d e v i c e d o e s not affect the s t a t u s of the s e r o t y p e d e s i g n a tions-they s t i l l a r e i n f r a s u b s p e c i f i c d e s i g n a t i o n s , equivalent
to, a n d r e p r e s e n t i n g , a n t i g e n i c f o r m u l a e . It m a y b e s u g g e s t e d that a u t h o r s m i g h t s p e l l out the c o m p l e t e n a m e of
e a c h S a l m o n e l l a s e r o t y p e t h e first t i m e that i t o c c u r s i n a
publication.
Subsequent r e f e r e n c e s to the s e r o t y p e s could
b e m a d e s i m p l y by using s e r o t y p e t y p h i m u r i u m , s e r . a n a t u m ,
o r by c a p i t a l i z a t i o n of t h e i n i t i a l l e t t e r . A c c o r d i n g to the
p r o p o s a l m a d e h e r e i n the so-called " c l a s s i c a l s p e c i e s ' '
should b e w r i t t e n i n t h e s a m e m a n n e r a s given i n t h e a b o v e m e n t i o n e d e x a m p l e s , e . g. S a l m o n e l l a e n t e r i d i t i s s e r o t y p e
e n t e r i d i t i s . The s p e c i f i c e p i t h e t s of t h e " c l a s s i c a l s p e c i e s , "
o t h e r than 2. c h o l e r a e - s u i s , 2. typhi a n d S. e n t e r i d i t i s should
b e r e j e c t e d a s s p e c i e s n a m e s but r e t a i n e d a s s e r o t y p e d e s i g n a t i o n s . Thus, the " c l a s s i c a l s p e c i e s " a l l m a y b e i n t e g r a t e d
e a s i l y into a single n o m e n c l a t u r a l , a s well a s t a x o n o m i c ,
s y s t e m by u s i n g the " c l a s s i c a l " e p i t h e t s a s s e r o t y p e d e s i g nations.
In o u r opinion, the m o s t i m p o r t a n t r e q u i r e m e n t s f o r
p r o g r e s s i n t h i s a r e a a r e t h e a c c e p t a n c e a n d u s e of a s i n g l e
s y s t e m of n o m e n c l a t u r e a n d t h a t w o r k e r s i n a l l f i e l d s u s e
that s y s t e m c o r r e c t l y .
If leading i n v e s t i g a t o r s i n b a c t e r i a l
g e n e t i c s a n d physiology c a n t a k e t h e t i m e a n d t r o u b l e to
l e a r n t h e m e a n s f o r a c c u r a t e c h a r a c t e r i z a t i o n of t h e s e b a c t e r i a and t o a t t e n d t o t h e d e t a i l s of c o r r e c t t e r m i n o l o g y , a s
s o m e have done, o t h e r s m i g h t follow t h e e x a m p l e .
8h. Although t h e c o r r e c t n a m e f o r t h e t h i r d s p e c i e s of
the genus S a l m o n e l l a , a s p r o p o s e d h e r e i n , a p p a r e n t l y i s
S. e n t e r i d i t i s ( G a e r t n e r ) C a s t e l l a n i and C h a l m e r s , it i s t h e
opinion of the a u t h o r t h a t a n exception t o t h e r u l e s m i g h t b e
i n o r d e r i n t h i s i n s t a n c e . Since i t h a s not b e e n e m p l o y e d
p r e v i o u s l y i n connection with s a l m o n e l l a e , t h e n e u t r a l n a m e
S a l m o n e l l a e n t e r i c a m i g h t b e c o n s e r v e d f o r u s e i n s t e a d of
S. e n t e r i d i t i s . In t h i s event, s u g g e s t e d u s a g e would b e the
s a m e a s that outlined i n the f o r e g o i n g p a r a g r a p h s , e . g . ,
S a l m o ne 1la e.nt e r i ca s e r 0typ e e nt e r i d i t i s , Sa lmon e 1la e n t e r i c a
(typhimurium) etc.
P r e c e d e n t h a s been e s t a b l i s h e d i n connection with t h e
r e j e c t i o n of " c l a s s i c a l s p e c i e s ' ' i n f a v o r of a l i m i t e d n u m b e r
of s p e c i e s , a s w e l l a s i n t h e u s e of i n f r a s u b s p e c i f i c d e s i g n a t i o n s f o r s e r o t y p e s within a s p e c i e s .
In s o m e i n s t a n c e s
t h e s e i n f r a s u b s p e c i f i c d e s i g n a t i o n s have r e p l a c e d t h e n a m e s
of l e g i t i m a t e l y published " c l a s s i c a l s p e c i e s . " F o r e x a m p l e ,
the g e n u s Shigella now i s l i m i t e d to f o u r s p e c i e s , which a r e
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f u r t h e r subdivided into s e r o t y p e s d e s i g n a t e d by A r a b i c n u m e r a l s . These n u m e r a l s a r e infrasubspecific designations,
e a c h of which i n d i c a t e s the a n t i g e n i c constitution of a s e r o type.
Thus, t h e A r a b i c n u m b e r s u s e d i n connection with
s h i g e l l a e a r e c o m p l e t e l y a n a l a g o u s to the i n f r a s u b s p e c i f i c
n a m e s given t o s e r o t y p e s of S a l m o n e l l a . Such " c l a s s i c a l
s p e c i e s " a s Shigella a m b i g u a (Shigella s c h m i t z i i ) a n d Shigella
n e w c a s t l e w e r e r e j e c t e d , a t l e a s t by i n f e r e n c e , with ' t h e
adoption of the p r e s e n t n o m e n c l a t u r a l s y s t e m , i n which t h e
f o r m e r b e c a m e Shigella d y s e n t e r i a e s e r o t y p e 2 a n d the l a t t e r
b e c a m e Shigella f l e x n e r i s e r o t y p e 6.
9. The g e n e r i c n a m e A r i z o n a a n d the s p e c i f i c epithet
a r i z o n a e w e r e given by Kauffmann a n d E d w a r d s i n 1952
(I.B.2:2).
9a. The f i r s t m e m b e r of the g r o u p p r e s e n t l y known a s
Arizona was isolated, described, and named Salmonella sp.
( D a r - e s - s a l a a m type v a r i e t y f r o m A r i z o n a ) ( s i c ) b y C a l d w e l l
a n d R y e r s o n (J. Infect. D i s e a s e s , e : 2 4 2 , 1939). A c c o r d i n g
to P r o f e s s o r Kauffmann's m o r e r e c e n t p r o p o s a l s (Acta P a t h .
M i c r o b i o l . Scand. 49:393, 1960; Kauffmann, l961), t h e A r i zona g r o u p m i g h t b e d e s i g n a t e d S a l m o n e l l a subgenus 111. If
s u c h a taxonomic change w e r e m a d e , t h e - A
group
m i g h t b e added to t h e g e n u s S a l m o n e l l a a s a f o u r t h s p e c i e s ,
S a l m o n e l l a a r i z o n a e Kauffmann (1 941) e m e n d .
10: The g e n e r i c t e r m C i t r o b a c t e r w a s p r o p o s e d by W e r k m a n a n d G i l l e n (J. B a c t e r i o l . 23:165, 1932) who t r a n s f e r r e d
t h e b a c t e r i u m d e s c r i b e d by B r a a k t o t h e genus a s C i t r o b a c t e r freundii.
11. A b r i e f r e v i e w of t h e n o m e n c l a t u r e of t h e t r i b e
K l e b s i e l l e a e w a s given by Ewing, Davis, a n d Johnson (I.B.
12:47, 1962). S i n c e the g e n e r i c t e r m K l e b s i e l l a T r e v i s a n
w a s c o n s e r v e d (Code, 166) a n d K l e b s i e l l a pneumoniae
( S c h r o e t e r ) T r e v i s a n w a s d e s i g n a t e d type s p e c i e s of the
c o n s e r v e d genus (Code, 166) i t i s c l e a r t h a t t h e n a m e of t h i s
t r i b e should b e K l e b s i e l l e a e .
1 2 . A b r i e f r e v i e w of the l i t e r a t u r e r e g a r d i n g t h e t e r m
"oxytoca" is i n o r d e r s i n c e the t e r m h a s b e e n u s e d i n a
v a r i e t y of combinations a n d i n connection with s e v e r a l b i o t y p e s of b a c t e r i a . B a c i l l u s oxytocus p e r n i c i o s u s (Wyssokow i t s c h ) F l u e g g e w a s i s o l a t e d f r o m s t a l e m i l k by Wyssokow i t s c h i n P r o f e s s o r F l u e g g e ' s l a b o r a t o r y . I t w a s n a m e d by
F l u e g g e a n d w a s d e s c r i b e d a s being gelatin n e g a t i v e . T r e v i s a n (1889, 1 7 , v. I . B . 2 : 1 3 , 1952) r e f e r r e d t o the above-
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m e n t i o n e d d e s c r i p t i o n a n d u s e d t h e n a m e B a c i l l u s oxytocus,
which w a s t h e f i r s t c o r r e c t b i n a r y combination u s e d i n connection with t h e m i c r o o r g a n i s m (validly p u b l i s h e d ) . M a c Conkey (J. Hyg. 6-:384, 1906) obtained a c u l t u r e of B,. 2t o c u s p e r n i c i o s u s f r o m t h e K r S l collection.
This culture,
which w a s supposedly f r o m F l u e g g e , w a s nonmotile, f a i l e d
t o f o r m indol, f a i l e d t o liquefy gelatin, a n d gave a p o s i t i v e
Voges-Proskauer test.
T h e s e r e a c t i o n s , a s well a s o t h e r s
l i s t e d by MacConkey, a r e s i m i l a r to t h o s e given by Klebs i e l l a pneumoniae s t r a i n s . The epithet w a s a l s o e m p l o y e d
by C a s t e l l a n i a n d C h a l m e r s (1919), who d e s c r i b e d a n i n d o l
p o s i t i v e b a c t e r i u m o t h e r w i s e like g . pneumoniae u n d e r the
d e s i g n a t i o n E s c h e r i c h i a o x y t o c u s . In 1909, MacConkey d e s c r i b e d gelatin p o s i t i v e , indole p o s i t i v e , nonmotile f o r m s
u n d e r t h e d e s i g n a t i o n B,. oxytocus p e r n i c i o s u s ( J . Hyg. 9:86,
1909). The t e r m A e r o b a c t e r oxytocum w a s e m p l o y e d i n
B e r g e y ' s Manual (1923) by Weldin (1927), a n d i n 4 t h edition
of B e r g e y ' s Manual f o r nonmotile, indol p o s i t i v e , g e l a t i n
negative b a c t e r i a having t h e g e n e r a l c h a r a c t e r i s t i c s of k l e b s i e l l a e . Thus i t i s c l e a r t h a t the m o s t authentic d e s c r i p t i o n
of B,. oxytocus p e r n i c i o s u s i s t h e c h a r a c t e r i z a t i o n of t h e
i s o l a n t f r o m t h e K r S l c o l l e c t i o n given by MacConkey (1906
loc. c i t . ) .
It a l s o i s a p p a r e n t that the epithet oxytocus i s a
l a t e r synonym of the e p i t h e t pneumoniae T r e v i s a n , 1885,
and h e n c e i l l e g i t i m a t e .
More recently s e v e r a l authors ( e . g. Lautrop, Acta P a t h .
M i c r o b i o l . Scand. 39:375, 1956) h a v e included indol p o s i t i v e ,
gelatin p o s i t i v e , s t r a i n s u n d e r the d e s i g n a t i o n "oxytoca," a s
did MacConkey (1909 ;
Loc. c i t . ) , a n d o t h e r s (Cowan e t 51.
J . G e n . M i c r o b i o l . Q:601, 1960) have doubted t h a t s u c h
c u l t u r e s should b e c a l l e d k l e b s i e l l a e .
The only e s s e n t i a l
d i f f e r e n c e between t h e oxytoca s t r a i n s a n d o t h e r indol p o s i tive kleb s i e l l a e p r e s e n t l y known i s g e l a t i n liquefaction, which
i s d e l a y e d . Hence, t h e r e i s l i t t l e doubt but that the b a c t e r i a
a r e Klebsiella.
However, i t i s b e l i e v e d t h a t the oxytoca
s t r a i n s a r e not sufficiently d i f f e r e n t to w a r r a n t s t a t u s a s a
separate species.
They a r e r e g a r d e d a s a biotype of 5 .
pneumoniae, which m a y b e c a l l e d the oxytoca biotype i f d e sired.
T h i s biotype would include c u l t u r e s t h a t a r e both
indol and g e l a t i n p o s i t i v e .
The s p e c i e s n a m e 5 . oxytoca should b e r e j e c t e d .
1 3 . Action by the J u d i c i a l C o m m i s s i o n i s pending o n
A e r o b a c t e r v s . Enterobacter (Hormaeche and Edwards, I.B.
10:71, 1960).
-
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1 4 . The b a c t e r i a c l a s s i f i e d i n the Hafnia group a r e
c l o s e l y r e l a t e d to A e r o b a c t e r a e r o g e n e s and a r e r e g a r d e d
a s a s u b s p e c i e s ( v a r i e t y ) of that s p e c i e s . It i s c l e a r t h a t
sufficient d i f f e r e n c e s e x i s t between m e m b e r s of t h e Hafnia
a e r o g e n e s to w a r r a n t the s u b g r o u p a n d c u l t u r e s of
s p e c i f i c d e s i g n a t i o n s . If additional d i f f e r e n c e s of taxonomic
i m p o r t a n c e a r e found i n f u t u r e , t h i s g r o u p of b a c t e r i a m a y
b e e l e v a t e d to s p e c i f i c r a n k , i n which c a s e t h e a p p r o p r i a t e
n a m e would b e A e r o b a c t e r hafniae. C o n v e r s e l y the g r o u p
m a y b e l o w e r e d to biotype s t a t u s i f f u t u r e s t u d i e s i n d i c a t e
However, it i s c l e a r t h a t g e n e r i c
that this is desirable.
r a n k i s not w a r r a n t e d a t t h i s t i m e .
1 5 . A e r o b a c t e r liquefaciens G r i m e s and H e n n e r t y (Sci.
P r o c . Roy. Dublin SOC. 20(NS):89, 1931) v s . A e r o b a c t e r
lipolyticus G r i m e s ( I . B . -=:111, 1961). The s p e c i e s n a m e
A e r o b a c t e r Liquefaciens B e i j e r i n c k i s r e g a r d e d a s i l l e g i t i m a t e s i n c e i t i s a l a t e r synonym of A e r o m o n a s hydrophila
(Chester) Stanier.
The b a c t e r i u m d e s c r i b e d b y B e i j e r i n c k
u n d e r the n a m e A e r o b a c t e r liquefaciens i s c o n s i d e r e d to b e
A e r o m o n a s hydrophila b y the a u t h o r
Ewing et ~ 1 1961).
.
Thus, i t would s e e m t h a t the e p i t h e t liquefaciens i s a v a i l a b l e
(Rule 26, Code, 97) f o r u s e , t h a t the combination A e r o b a c t e r
l i q u e f a c i e n s G r i m e s and H e n n e r t y i s l e g i t i m a t e , and t h a t
&. lipolyticus i s a l a t e r synonym.
However, a r u l i n g on lipolyticus v s . liquefaciens a n d
c o n s e r v a t i o n of 4. liquefaciens o r
Qolyticus i s d e s i r a b l e .
16. The n o m e n c l a t u r e a n d taxonomy of the genus S e r r a t i a
w e r e d i s c u s s e d by Ewing, Davis, and J o h n s o n ( I . B . z : 4 7 ,
1962).
1 7 . The n o m e n c l a t u r e a n d taxonomy of the t r i b e P r o t e e a e
w e r e d i s c u s s e d by Ewing (I.B. 5 : 1 0 2 , 1962).
A.
(v.
A.
A s s t a t e d a t t h e o u t s e t , t h e outline of n o m e n c l a t u r e f o r
the f a m i l y E n t e r o b a c t e r i a c e a e p r e s e n t e d follows the t a x o n o m i c s y s t e m published e a r l i e r (Ewing a n d E d w a r d s 1960,
r e v i s e d 1962) a n d the b i o c h e m i c a l m e t h o d s given i n t h a t
publication m a y b e r e g a r d e d a s t h e b a s i s f o r the t r i b e s ,
g e n e r a , a n d s p e c i e s given i n t h e outline of n o m e n c l a t u r e .
The t e r m i n o l o g y e m p l o y e d i n t h e outline i s n o m e n c l a t u r a l l y l e g i t i m a t e , a s f a r a s we have b e e n a b l e to d e t e r m i n e .
However, t h e r e a r e a n u m b e r of points on which a c t i o n by
the Judicial Commission i s e i t h e r n e c e s s a r y o r desirable.
T h e s e a r e s u m m a r i z e d a s follows, f o r convenience:
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AND TAXONOMY
1 . S i n c e the type s p e c i e s of the genus E s c h e r i c h i a w a s
d e s i g n a t e d a s E s c h e r i c h i a &i,
it m a y b e i n f e r r e d t h a t the
s p e c i f i c epithet &i w a s c o n s e r v e d . However, definite cons e r v a t i o n of E s c h e r i c h i a
would b e d e s i r a b l e . (Annot. 3 ) .
2. The s p e c i e s n a m e S a l m o n e l l a typhi ( o r 2. t y p h o s a )
should b e c o n s e r v e d . The n a m e that i s not c o n s e r v e d should
b e r e j e c t e d . (Annot. 7).
3 . The n a m e S a l m o n e l l a kauffmannii should b e r e j e c t e d
a s a n a m e f o r a t h i r d s p e c i e s of S a l m o n e l l a . (Annot. 8 a ) .
4. The n a m e S a l m o n e l l a e n t e r i d i t i s ( o r 2 . e n t e r i c a )
should be c o n s e r v e d f o r u s e a s the n a m e of the t h i r d s p e c i e s
of S a l m o n e l l a . (Annot. 8 a , 8h). If S a l m o n e l l a e n t e r i c a i s
d e e m e d u n a c c e p t a b l e , t h i s n a m e should b e r e j e c t e d a s far
a s u s a g e i n t h i s context i s c o n c e r n e d .
5. 'Arizona a r i z o n a e Kauffmann a n d E d w a r d s i s n a m e d
h e r e i n a s the type s p e c i e s of the genus A r i z o n a .
6. The s p e c i f i c n a m e K l e b s i e l l a oxytoca should b e r e j e c t e d , s i n c e i t i s a l a t e r objective synonym of K l e b s i e l l a
pneumoniae
(Annot. 1 2).
7 . Action should b e taken on t h e s u b j e c t of A e r o b a c t e r v s .
E n t e r o b a c t e r , with c o n s e r v a t i o n of one a n d r e j e c t i o n of the
o t h e r . (Annot. 1 3 ) .
8. C o n s e r v a t i o n of t h e s p e c i f i c n a m e A e r o b a c t e r liquef a c i e n s G r i m e s a n d H e n n e r t y o r A e r o b a c t e r lipolyticus
G r i m e s i s needed. (Annot. 15).
9 . P r o p o s a l s m a d e r e g a r d i n g the n o m e n c l a t u r e of t h e
genus S e r r a t i a a n d the t r i b e P r o t e e a e should be a c t e d on.
(Annot. 16, 1 7 ) .
e
.
The a t t e n t i o n of m e m b e r s of t h e J u d i c i a l C o m m i s s i o n a n d
of the N o m e n c l a t u r e C o m m i t t e e i s d i r e c t e d to t h e a b o v e m e n t i o n e d p o i n t s . It i s r e s p e c t f u l l y r e q u e s t e d t h a t t h e p o i n t s
b e c o n s i d e r e d and t h a t n e c e s s a r y a c t i o n b e taken on t h e m .
F u r t h e r , i t is respectfully requested that the proposed
n o m e n c l a t u r a l s y s t e m , a s given i n the outline, b e c o n s i d e r e d
a n d t h a t a c t i o n b e taken t o d e t e r m i n e w h e t h e r the s y s t e m i s
nomenclaturally c o r r e c t .
Whether o r not a s y s t e m of n o m e n c l a t u r e i s "accepted"
depends upon u s a g e .
The s y s t e m p r e s e n t e d h e r e i n w a s developed d i r e c t l y f r o m a p r a c t i c a l t a x o n o m i c s c h e m a which
h a s b e e n u s e d t o c o n s i d e r a b l e advantage f o r a n u m b e r of
y e a r s , a n d which h a s found r a t h e r wide a c c e p t a n c e . Hence,
i t would s e e m t o the w r i t e r t h a t a n o m e n c l a t u r a l s y s t e m
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d e r i v e d f r o m a t e s t e d p r a c t i c a l taxonomic s c h e m a should b e
u s e f u l a n d a d v a n t a g e o u s . The a u t h o r c a n s e e no r e a s o n why
the n o m e n c l a t u r e outlined h e r e i n cannot b e u s e d to advantage
by i n v e s t i g a t o r s i n b a c t e r i a l physiology, g e n e t i c s , o r a n y
o t h e r field, when they a r e working with m e m b e r s of the
family Enterobacteriaceae.
S t r a i n s of a p a r t i c u l a r s p e c i e s
t h a t d i f f e r f r o m t h e p a r e n t f r o m which they w e r e d e r i v e d ,
which p r o d u c e s o m e s u b s t a n c e o r f a i l t o d o so, o r which a r e
dependent upon o r r e s i s t a n t t o s o m e agent, m a y e a s i l y b e
d i s t i n g u i s h e d by the u s e of a s u i t a b l e s y m b o l along with t h e
c o r r e c t n o m e n c l a t u r a l d e s i g n a t i o n . U s a g e of c o r r e c t nomenc l a t u r e m a y b e e x p e c t e d to lend c l a r i t y t o t h e publications
of w o r k e r s i n a l l f i e l d s of b a c t e r i o l o g y . P a r e n t h e t i c a l l y , we
might note t h a t a d d i t i o n a l s p e c i e s o r s u b s p e c i e s m a y b e
a d d e d t o the v a r i o u s g e n e r a when the n e e d a r i s e s a n d a l s o
additional g e n e r a o r t r i b e s should it b e c o m e n e c e s s a r y .
However, additions should not b e m a d e i n d i s c r i m i n a t e l y .
A s s t a t e d e a r l i e r , t h e w r i t e r i s of the opinion t h a t the
m o s t i m p o r t a n t r e q u i r e m e n t s i n t h i s a r e a a r e the adoption
of a single s y s t e m of n o m e n c l a t u r e f o r t h e f a m i l y E n t e r o b a c t e r i a c e a e a n d t h a t i n v e s t i g a t o r s i n a l l f i e l d s of b a c t e r i o l ogy u s e t h a t s y s t e m c o r r e c t l y when dealing with m e m b e r s
of t h e f a m i l y . We b e l i e v e t h a t t h e n o m e n c l a t u r e outlined
h e r e i n c a n b e r e c o m m e n d e d a s s u c h a single s y s t e m , t h a t
i t s u s e w i l l p r o v e to b e advantageous, a n d that if u s e d , it
eventually w i l l find a c c e p t a n c e .
Additional R e f e r e n c e s
B e r g e y , D. H. 1923. Manual of D e t e r m i n a t i v e B a c t e r i o l o g y .
W i l l i a m s a n d Wilkins, B a l t i m o r e .
B r e e d , R. S . , E.G. D. M u r r a y , a n d A. P. H i t c h e n s . 1928.
B e r g e y ' s Manual of D e t e r m i n a t i v e B a c t e r i o l o g y . 7th e d .
W i l l i a m s a n d Wilkins, B a l t i m o r e .
-I -a n d N . R . S m i t h . 1 9 4 7 . B e r g e y ' s Manual of
Determinative Bacteriology.
7th e d . W i l l i a m s and
Wilkins, B a l t i m o r e .
C a s t e l l a n i , A . a n d A . J . C h a l m e r s . 1919. A Manual of
T r o p i c a l Medicine, 3 r d e d . W i l l i a m Wood a n d Company,
New Y o r k .
and
1920. S u r la c l a s s i f i c a t i o n de c e r t a i n s
g r o u p s de b a c i l l e s a 6 r o b i e s d e l ' i n t e s t i n e humain. Ann.
Inst. Pasteur 2 : 6 0 0 - 6 2 1
.
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AND TAXONOMY
E d w a r d s , P . R . a n d W . H . Ewing. 1962. Identification of
E n t e r o b a c t e r i a c e a e . 2nd e d . B u r g e s s , Minneapolis.
Ewing, W . H . , I . S u a s s u n a , a n d 1 . R . S u a s s u n a . 1960. The
b i o c h e m i c a l r e a c t i o n s of m e m b e r s of the P r o t e u s g r o u p .
CDC publication. C o m m u n i c a b l e D i s e a s e C e n t e r ,
Atlanta.
, R . Hugh a n d J . G . J o h n s o n . 1961. Studies on the
A e r o m o n a s g r o u p . CDC publication. C o m m u n i c a b l e
Disease Center, Atlanta.
a n d P . R . E d w a r d s . 1962. T h e p r i n c i p l e d i v i s i o n s
a n d g r o u p s of E n t e r o b a c t e r i a c e a e . ( R e v i s e d f r o m I n t e r n a t l . B u l l . B a c t . Nomen. T a x . 2 : 1 - 1 2 . 1960). CDC
publication. C o m m u n i c a b l e D i s e a s e C e n t e r , A t l a n t a .
Kauffmann, F. 1941. Die B a k t e r i o l o g i e d e r S a l m o n e l l a G r u p p e . E i n a r M u n k s g a a r d , Copenhagen.
1949. On t h e c l a s s i f i c a t i o n of E n t e r o b a c t e r i a c e a e .
A c t a P a t h . Microbiol. Scand. G : 8 7 9 - 8 8 1 .
1953. On the c l a s s i f i c a t i o n a n d n o m e n c l a t u r e of
E n t e r o b a c t e r i a c e a e . Riv. I n s t . S i e r o t e r a p . I t a l . 28:485491.
1956. Z u r b i o c h e m i s c h e n und s e r o l o g i s c h e n G r u p p e n
-und Typen-Enteilung d e r E n t e r o b a c t e r i a c e a e .
C e n t r a l . f . B a k t . , I O r i g . 165:344-354.
1961. Die B a k t e r i o l o g i e d e r S a l m o n e l l a - S p e c i e s .
Munksgaard, Copenhagen.
S u a s s u n a , I. a n d I . R . S u a s s u n a . 1960. S o b r e p o s i c a o s i s tematica de Proteus americanus Pacheco, Proteus paraa m e r i c a n u s Magalhaes and Aragao E., and Eberthella
a l c a l i f a c i e n s De S a l l e s G o m e s . A n a i s de m i c r o b i o l .
8:161-168.
Weldin, J . O . 1927. The colon-typhoid group of b a c t e r i a a n d
r e l a t e d f o r m s . R e l a t i o n s h i p s a n d c l a s s i f i c a t i o n . Iowa
State Colt. J o u r . Sci. 1:121-197.
.
.
.
.
ADDENDUM
A f t e r t h i s p a p e r w a s r e c e i v e d , the J u d i c i a l C o m m i s s i o n
of t h e I n t e r n a t i o n a l N o m e n c l a t u r e C o m m i t t e e a c t e d upon the
r e q u e s t of H o r m a e c h e a n d E d w a r d s (I.B. s : 7 7 - 7 8 , 1960)
and c o n s e r v e d t h e g e n e r i c n a m e E n t e r o b a c t e r H o r m a e c h e
a n d E d w a r d s (Opinion 28, I.B. g : 3 8 , 1963). T h u s , G e n u s 11
of T r i b e 111 b e c o m e s :
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BULLETIN
G e n u s I1 E n t e r o b a c t e r H o r m a e c h e a n d E d w a r d s
1 . E n t e r o b a c t e r c l o a c a e ( J o r d a n ) H o r m a e c h e and E d w a r d s
2 . Enterobacter a e r o g e n e s (Kruse) Hormaeche and
Edwards
2a. E n t e r o b a c t e r a e r o g e n e s s u b s p e c i e s hafniae
( M o e l l e r ) Ewing comb. nov.
3 . E n t e r o b a c t e r liquefaciens ( G r i m e s and H e n n e r t y )
Ewing c o m b . nov.
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