INTERNATIONAL BULLETIN O F BACTERIOLOGICAL
NOMENCLATURE AND TAXONOMY
Volume 6
No. 3
July 15, 1956
pp. 121-128
THE CLASSIFICATION O F LOWER ORGANISMS
Nom enclatur a1 Review
.
The Classification of Lower
Copeland, H e r b e r t Faulkner
O r g a n i s m s . P a c i f i c Books. P a l o Alto, California. pp. ix,
302. 1956
The Linnaean assumption that all living things m a y be
allocated e i t h e r to the plant kingdom o r to the a n i m a l kingdom h a s been challenged on a number of o c c a s i o n s . P r o p o s a l s w e r e e a r l y made that the relatively s i m p l e f o r m s ,
including the P r o t o z o a and Protophyta, might well be c o m bined into a t h i r d kingdom distinct f r o m P l a n t a e and Anim a l i a . Among the names u s e d have been P r o t o c t i s t a ( H G ,
1860), P r i m a l i a (Wilson and C z s s i n , 1864) and P r o t i s t a
( H a e c k e m . Copeland a g r e e s with c e r t a i n r e c e n t w r i t e r s t h a t four kingdoms should be recognized.
The author s t a t e s that "the purpose of this work is to
p e r s u a d e the community of biologists that the accepted p r i m a r y classification of living things a s two kingdoms, plants
and a n i m a l s , should be abandoned; that the kingdoms of
plants and a n i m a l s a r e to be given definite l i m i t s , and that
the o r g a n i s m s excluded f r o m them a r e to be organized as
two other kingdoms, I ' He n a m e s and defines the four kingdoms as follows:
-
Kingdom I. Mychota Enderlein, 1925. (Protophyta Haeckel,
1894; Monera Copeland, 1938; Anucleobionta R o t h m a l e r ,
1948). O r g a n i s m s without nuclei; the b a c t e r i a and the
blue-green algae.
Kingdom 11. P r o t o c t i s t a Hogg, 1860. P r o t o z o a Haeckel a
1894; P r o t i s t a Copeland, 1938; Protobionta Rothmaler,
1948). Nucleate o r g a n i s m s not of the c h a r a c t e r s of plants
and animals; the protozoa, the r e d and brown algae and
the fungi.
Kingdom 111. P l a n t a e (Metaphyta Haeckel, 1894; Cormobionta Rothmaler, 1948). O r g a n i s m s i n whose c e l l s o c c u r
c h l o r o p l a s t s , being plastids of a b r i g h t g r e e n c o l o r , containing the pigments chlorophyll 2, chlorophyll k, c a r o tin and xanthophyll, and no o t h e r s ; and which produce
s u c r o s e , t r u e s t a r c h , and t r u e cellulose.
Kingdom IV. Animalia. (Metazoa Haeckel, 1894; G a s t r o bionta Rothmaler, 1948). Multicellular o r g a n i s m s which
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pass during development through the stages called blastula and gastrula; typically predatory, and accordingly
consisting of unwalled cells and attaining high complexity
of structure and function.
We a r e concerned in this review only with the kingdom
Mychota defined to include the bacteria and the blue-green
algae. As noted above, the distinctive character ascribed'
to the members of this kingdom i s the absence of a nucleus.
The nucleus i s defined as follows:
"Morphologically the nucleus is a p a r t of a protoplast
which i s s e t apart by a membrane and which originates
ordinarily by a division of a pre-existing nucleus in
the manner called mitosis. In this process, a definite
number of definite chromosomes appear and undergo
equal division. I t
The author senses the difficulty in an unqualified statement
that these organisms lack nuclei. He says:
"In describing the Mychota as lacking nuclei, one commits himself to one side of a controversy of many y e a r s
duration. Because of the greater size of the cells of the
blue-green algae, the facts a r e more easily ascertained
in this group than in the proper bacteria."
The latest cited authority on the cytology of the blue-green
algae is Haupt (1923), who emphasized "the impropriety of
calling any p a r t of these cells a nucleus. It Copeland states
further, "Recent studies of typical bacteria by conventional
microtechnical methods and by the electron microscope have
made i t possible to recognize the essential identity of the
structure of their cells with those of the blue-green algae. I '
To the writer the proof of the presence o r absence of
a nuclear membrane as a criterion of the presence of a
nucleus is rather beside the point. Are there bodies within
the cells of the bacteria which perform the essential functions of chromosomes in the mechanism of heredity?
The
evidence of recent y e a r s has accumulated to the pointwhere
there i s little remaining doubt the answer is yes, and that
withinthe bacterial cell there i m a bodywhich functions much
a s does the nucleus of a higher plant o r animal. The author
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BACTERIOLOGICAL NOMENCLATURE
AND T A X O N O MY
tacitly admits the existence of this fundamental difficulty in
his scheme of classification. The evidence certainly does
not warrant the separation of the bacteria from other plants
on the basis of the postulated lack of a nucleus i n the cell.
It would seem that the intergradations between the bluegreen algae and the bacteria would warrant the assumption
of interrelationship, but with no good reason to place them
in a separate kingdom. Thimann (1955), in his text on microbiology states the c a s e a s follows:
"Whether o r not bacteria possess a nucleus has been a
subject of discussion and argument for forty y e a r s and
has only been settled (in the affirmative) relatively
recently.
Also, "Taking the observations as a whole,
there can be no doubt that bacteria possess one o r two
nuclear granules or chromosomes.
Notwithstanding disagreement with Copeland's thesis,
there is much of nomenclatural and taxonomic interest in
his discussion of the Mychota. He writes:
"It is the right and duty of every person who thinks that
the taxonomic system can be improved to propose to
change it. A salutary convention requires that proposals i n taxonomy be unequivocal; one proposes a change
by publishing it as in effect; i t comes actually into
effect in the degree that the generality of students of
classification accept i t . I '
The author follows Oparin (1938) in his concept of the
situation on the e a r t h when life began, in water saturated
with the gases ammonia and methane of which the atmosphere presumably consisted. "These theories indicate, as
the most primitive f o r m of life which has been able to s u r How anaerobes could secure
vive, the anaerobic bacteria.
their growth energy from the components of the primitive
atmosphere as specified i s not stated. The conclusion i s
reached "Life requires energy" and "Under anaerobic conditions, an organism can obtain energy by converting sugar
to alcohol, but it cannot u s e alcohol as a source of energy. ' I
In fact, however, there a r e bacteria that, under anaerobic
conditions i n a suitable environment, can utilize alcohol as
a source of energy.
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"The existence of organisms without nuclei shows that
the nucleus evolved after life did; i t did not evolve at the
same time as the protoplasm.
If the modern work on the
viruses is to be trusted i t would seem quite as probable
that nuclear material and the essential components of the
nucleus evolved at least a s soon a s the protoplasm.
Copeland's Chapter I1 i s "An Essay on Nomenclature.
That there a r e three Codes of Nomenclature, Botanical,
Zoological, and Bacteriological, i s pointed out. Quite appropriately i t is emphasized that the very existence of three
s e t s of rules and their continual amendment i s "evidence of
imperfection." He then points out "certain anomalies in the
codes as they stand." The Zoological Rules t e r m the second
component of the name of a species a specific name. Copelant insists that the principles of grammar a r e violated by
"treating specific epithets a s names .ll The sapiens of Homo
sapiens is not a name of man, nor i s the adjective vulgare
in Hordeum vulgare the name of barley. The name of a
species i s a binary combination. Botanists and bacteriologists use the phrases "species name" and "specific name"
synonymously, and designate the two components the gene r i c name and the specific epithet, respectively.
The author also calls attention to the fact that in zoology
all family names end in -idae and all subfamily names in
-inae. These a r e plural feminine adjectival endings, and
should agree with the name of the kingdom. Animalia i s a
plural neuter noun. He suggests that these endings were
f i r s t used f o r the families of the birds (Aves, fern. pl.),
and the family names a r e unobjectionable in this group, but
quite inappropriate in a kingdom Animalia.
He criticizes the Botanical Code in that i t enunciates a
s e r i e s of principles followed by a l i s t of rules which in
some c a s e s prescribe procedures not warranted by the
preceding principles. He regards the l i s t of conserved and
rejected names (covering about 7 0 pages) as fixed quite
irrespective of principle. He Concludes that current botanical nomenclature "is to a considerable extent governed by
enactments of the nature of ex post facto laws and bills of
attainder.
The author very neatly points up the difficulty of getting
nomenclatural cooperation with zoologists. The Bacteriological Code provides that a bacterial generic name is illegitimate if it i s a later homonym of the name of a genus of
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AND TAXONOMY
the protozoa. Copeland notes that this is a u n i l a t e r a l a r rangement: "The avoidance of homonyms which they d e s i r e
will not, however, be attained; no zoologist will allow a new
name f o r the flagellate Klebsiella on account of an e a r l i e r
Klebsiella among b a c t e r i a . " This is probably t r u e a t p r e s ent. The g r e a t tragedy of biological nomenclature i s that
the botanists and zoologists have made no r e a l effort to
harmonize the wholly indefensible incompatibilities of t h e i r
p r e s e n t codes. Why should the e d i t o r s and u s e r s of Biolog i c a l A b s t r a c t s be compelled to obey t h r e e different codes,
s o m e t i m e s all t h r e e i n a review of a single a r t i c l e ? The
differences i n the codes a r e usually not g r e a t , but they a r e
s o m e t i m e s highly annoying.
Copeland r e a s o n s that by formation of two new kingdoms
I I a nomenclature
of s u p e r i o r legitimacy c a n be applied i n
groups t r e a t e d as r e m o v e d f r o m jurisdiction of the r u l e s .
He then proceeds to 'la brief formulationof those p r i n c i p l e s ,
appealing t o r e a s o n and proven sound i n p r a c t i c e , t o which
all nomenclature m u s t conform.
The f i r s t of t h e s e principles (not differing i n e s s e n t i a l s
f r o m that of e a c h of the t h r e e nomenclatural c o d e s ) i s s p e l l e d
out i n r e f r e s h i n g detail.
"Scientific names a r e w o r d s of the Latin language. They
a r e not 'of Latin f o r m ' o r 'construed as Latin;' they a r e
Latin. This is to t r e a t Latin as a living language and
scientific n a m e s as subject to the r u l e s of i t s g r a m m a r . "
Altogether seven principles a r e l a i d down, none of them
differing e s s e n t i a l l y f r o m those found i n all t h r e e c o d e s .
One s t a t e m e n t is well worth considering. It i s not found
i n any code at p r e s e n t but could advantageously be included
i n t h a t of the b a c t e r i o l o g i s t s . It r e a d s :
" I t follows f r o m the principle of the binomial nomenc l a t u r e of s p e c i e s that no genus is named until one o r
m o r e of its s p e c i e s a r e designated by binomial n a m e s .
The r e v i e w e r would go a s t e p f u r t h e r and make c l e a r that a
g e n e r i c name given by a n author without designation o r i n dication of the n a m e s of one o r m o r e contained s p e c i e s ,
even though validly published, is i l l r gitimate, and should so
r e m a i n u n t i l a s p e c i e s i n the genus h a s been n a m e d by a
subsequent author.
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C opeland's p r e s c r i p t i o n f o r capitalization of specific
epithets a c c o r d s with the e a r l i e r Botanical Codes, but with
none of the r e c e n t revisions of any of the codes.
The discussion of the u s e of p l u r a l s of g e n e r i c n a m e s is
internally inconsistent. The author s t a t e s " P l u r a l s of gene r i c names a r e not tenable" and then p r o c e e d s to show how
they may be u s e d p r o p e r l y . " E r i c a e m e a n s the s p e c i e s of
the genus E r i c a . ' I He points o u t , quite c o r r e c t l y , that such
a p l u r a l should n e v e r "be u s e d to designate the genus together with its a l l i e s . ' '
However, the author g e t s on untenable ground when he
says: "Names consisting of words other than g e n e r i c n a m e s
modified by terminations signifying 'resembling' o r 'of the
group of' a r e pot tenable, because they a r e nonsense. 'I He
i l l u s t r a t e s by u s e of E n g l e r ' s name Coniferinae applied to
a c l a s s objectionable because the Latin word conifer is an
adjective, and Coniferinae would p r e s u m a b l y m e a n " r e s e m bling cone-bearing." But t h e r e a r e many Latin nouns which
end i n f e r . A s t a n d a r d - b e a r e r is a signifer o r an aquilifer,
while vexillifer ( s t a n d a r d - b e a r i n g ) is a n adjective. Conifer
i n modern Latin c a n b e a noun. c o n e - b e a r e r . and Coniferinae m e a n "those resembling- c o n e - b e a r e r s . "
Copeland analyzes c o r r e c t l y the significance of - a c e a e
as a family ending . i n the kingdom P l a n t a e . Such family
n a m e s a r e Ifadjectives i n the feminine, agreeing with the
n a m e of the kingdom P l a n t a e . I' He i n s i s t s ( a p p r o p r i a t e l y )
that n a m e s of o r d e r s , f a m i l i e s , e t c . , of b a c t e r i a (usually
placed i n the kingdom P l a n t a e ) should, when placed i n the
kingdom Mychota, have the endings modified to indicate that
they a r e adjectives modifying a p l u r a l neuter noun
chota). Ordinal n a m e s should end i n -&a i n s t e a d of - a l e s ,
and family n a m e s i n - a c e a r a t h e r than - a c e a e . T o the r e viewer it s e e m s probable that the author is c o r r e c t i n s o m e
and i n e r r o r i n o t h e r s of the suggested changes. The Latin
endings -aceus, - a c e a , -aceum (singular); - a c e a s - a c e a e ,
- a c e a ( p l u r a l ) , give the feminine Latin p l u r a l - a c e a e (as i n
C o r y n e b a c t e r i a c e a e ) and the neuter p l u r a l i n - a c e a (Coryn e b a c t e r i a c e a ) . Copeland r e l a b e l s about 60 f a m i l i e s newly
'named o r t r a n s f e r r e d t o Mychota. He a l s o changes the ordinal ending -*s
to - a l e a i n t r a n s f e r r i n g f r o m P l a n t a e to
Mychota. The Latin endings -=s,
-=s,
-ale (aingular),
- a l e s , - a l e s , -&a
(plural), give feminine Latin p l u r a l s
w t i n o m y c e t a l e s ) and n e u t e r p l u r a l s i n -a&a (not
i n -a&s
-
'
(9-
-
-
-
-
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-*
-alea)(Actinomycetalia). The ending
c o m e s under the
L x n r u l e s o l declension of adjective s t e m s i n -i of two
endings. Five ordinal n a m e s should be c o r r e c t l y s p e l l e d
Ac tinomyce talia,
,Myxobacteralia, Sphaerotilalia, and Spirochaetalia. , T h e family n a m e s Achromob a c t e r a c e a , Azotobac t e r a c e a , Acetobac t e r acea, Nitrobac
t e r a c e a , Caulobacteracea, and Myxobacteracea a r e m i s spelled by Copeland, t h e r e is n o i i n the family ending - a c e a
and none i n the s t e m ending.
Five new f a m i l i e s a r e n a m e d and described: Kurthiacea,
P a s t e u r e l l a c e a , Chromatiacea, Rhodobacillacea, Chlorobiacea.
-
-
SUPRAGENERIC T A U IN THE KINGDOM MYCHOTA
I. Kingdom Mychota Enderlein.
A. Phylum Archezoa Haeckel.
1 . C l a s s Schizophyta (Cohn) McNab.
a . O r d e r Schizosporea Cohn. 13 F a m i l i e s .
Micrococcacea P r i b r a m ; N e i s s e r i a c e a P r g v o t ;
C o r y n e b a c t e r i a c e a L e h m a m and Neumann;
Rickettsiacea Pinkerton; Kurthiacea f a m . nov. ;
Bacillac e a Fis che r ; Ac hromobac t e r a c e a B r e e d ;
P a s t e u r e l l a c e a f a m . nov.; Rhizobiacea Conn;
Azotobacteracea Bergey, B r e e d and M u r r a y ;
Spirillac e a Migula; Ace tobac t e r ac e a B e r gey ,
B r e e d and Murray; N i t r o b a c t e r a c e a Buchanan.
b. O r d e r Ac tinomyc e t a l i a Buc hanan.
Mvcobacteriacea C h e s t e r .
C.
1 Family.
O r d e r Caulobacteralia Henrici and Johnson.
2 F a m i l i e s . Leptotrichacea S c h r o t e r :
Caulobacteracea Henrici and Johnson.
2 . C l a s s Mvxoschizomycetes Schaffner.
a. O r d e r Myxobacteralia C l e m e n t s . 5 F a m i l i e s .
Cytophagacea S t a n i e r ; Archangiacea Jahn;
Sorangiacea Jahn; Myxobacteracea E.F. Smith;
Myxococcacea Jahn.
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b. Order Spirochaetalia Buchanan. 2 Families.
Spir ochaetac ea Swellengr ebel;
Treponematacea Robinson
3. Class Archiplas tidea Bes s ey
.
a . Order Rhodobacteria Molisch. 3 Families.
Chromatiacea (Migula) nom. fam. nov. :
Rhodobacillac ea nom nov. ;
Chlorobiacea nom. nov.
.
b. Order Sphaerotilalia nom. nov.
Shpaerotilacea P r i b r a m .
1 Family.
I
C.
Order Cocco onea Campbell. 2 Families.
C h r ooc*ag
eli 1 R ab enho r s t ;
Achromatiacea Buc hanan.
d. Order Gloiophycea Rabenhorst. 10 Families.
Oscillatoracea Harvey; Beggiatoacea Migula;
Chamaesiphonacea Borzi; Nos tocacea (Nageli)
Rabenhorst; Scytonematacea Rabenhorst;
Chlamydotric hac ea P r i b r am; Rivulariac e a
Rabenhorst: SirosiDhonacea Rabenhorst:
Pleur oc aps ac e a Gei tle r ; C r enotric hac ea
Hansgirg.
Copeland ends his discussion: ''Such a r e the Mychota, the
organisms which may be characterized ae lacking nuclei. ' I
As i s evident from the above outline, Copeland'e arrangement of the bacteria differs quite materially from that of
most recent authors. F o r example, the inclusion of Rickettsiacea between genera a s unlike as Corynebacterium and
Bacillus among the true bacteria is quite in contrast to
recent tendency to emphasize their intermediate position
between the bacteria and viruses.
Apparently the s e v e r a l new taxa named and described
a r e validly published. Their incorporation into conventional
[bacteriological taxonomy will require some emendation of
spelling of suprageneric taxa.
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