International Journal of Systematic and Evolutionary Microbiology (2015), 65, 2769– 2773
DOI 10.1099/ijs.0.000336
Reclassification of Bacillus invictae as a later
heterotypic synonym of Bacillus altitudinis
Yang Liu, Qiliang Lai, Juan Du and Zongze Shao
Correspondence
Zongze Shao
[email protected]
State Key Laboratory Breeding Base of Marine Genetic Resources; Key Laboratory of Marine
Genetic Resources, Third Institute of Oceanography, SOA; Fujian Collaborative Innovation Center
for Exploitation and Utilization of Marine Biological Resources; Key Laboratory of Marine Genetic
Resources of Fujian Province, Xiamen 361005, PR China
The aim of this study was to reclarify the taxonomic status of strain Bacillus invictae Bi.FFUP1T by
performing comparative analyses with the other four type strains within the Bacillus pumilus
group. The digital DNA–DNA hybridization (dDDH) and average nucleotide identity (ANI) values
between strains B. invictae Bi.FFUP1 T (5DSMZ 26896T5MCCC 1A07089T), B. altitudinis
41KF2bT (5DSMZ 21631T5MCCC 1A06452T), B. safensis FO-36bT (5DSMZ
19292T5MCCC 1A6451T), B. pumilus ATCC 7061T (5DSMZ 27T5MCCC 1A06453T) and
B. xiamenensis HYC-10T (5MCCC 1A00008T) were, respectively, 82.90 % and 98.10 %,
which are greater than the thresholds for bacterial species delineation, suggesting that they
should belong to the same species, while the dDDH and ANI values between strain B. invictae
DSMZ 26896T and the other three type strains within the B. pumilus group were below the
respective thresholds of 70 % and 95 %. Meanwhile, B. invictae DSMZ 26896T and
B. altitudinis 41KF2bT shared 98.7 % gyrB gene sequence similarity based on resequencing,
whereas strain B. invictae DSMZ 26896T shared low similarities (,95 %) with the other three
type strains. In addition, in comparison with those from the other three type strains, phenotypic
data of B. invictae DSMZ 26896T and B. altitudinis 41KF2bT, including API 20NE, API ZYM,
Biolog GN2 and API 50CHB tests, showed slight differences. The data from these combined
genotypic and phenotypic analyses suggest that Bacillus invictae Branquinho et al. 2014
should be regarded as a later heterotypic synonym of Bacillus altitudinis Shivaji et al. 2006.
Bacillus invictae was isolated from a contaminated health
product in Portugal and then proposed, on the basis of
data from a polyphasic taxonomic approach, to represent
a novel species of the Bacillus pumilus group with a sole
strain (Branquinho et al., 2014a). Therefore, the B. pumilus
group comprises five species at the time of writing, including B. pumilus, B. safensis (Satomi et al., 2006), B. altitudinis (Shivaji et al., 2006), B. xiamenensis (Lai et al., 2014)
and B. invictae (Branquinho et al., 2014a). However,
these closely related species were not discriminated accurately on the basis of 16S rRNA gene sequence. Several
reliable and generally accepted typing methods, such as
single housekeeping gene sequence analysis (i.e. gyrB,
rpoB) (Branquinho et al., 2014b), multilocus sequence
analysis (Liu et al., 2013) and matrix-assisted laser desorption/ionization time-of-flight MS (Branquinho et al.,
Abbreviations: ANI, average nucleotide identity; dDDH; digital DNA–
DNA hybridization; DDH, DNA–DNA hybridization.
The GenBank/EMBL/DDBJ accession number for the draft genome
sequence of Bacillus invictae DSMZ 26896T is JXAI00000000.
000336 G 2015 IUMS
2014c), provide a promising avenue for identification of
strains within the B. pumilus group.
According to Branquinho et al. (2014a), two type strains,
B. invictae Bi.FFUP1T and B. altitudinis 41KF2bT, shared a
DNA–DNA hybridization (DDH) value of 61.7 % and
98.7 % gyrB gene sequence similarity. But, in the study of
Liu et al. (2013), the range of 95–96 % gyrB gene sequence
similarity was proposed as an interspecies gap for the
B. pumilus group. Therefore, in order to address this contradiction, it was necessary to redetermine the taxonomic
status of the type strain B. invictae Bi.FFUP1T using a variety
of genotypic and phenotypic criteria.
Four type strains, B. invictae DSMZ 26896T (5Bi.FFUP1T5
MCCC 1A07089T), B. altitudinis DSMZ 21631T (541KF2bT5
MCCC 1A06452T), B. safensis DSMZ 19292T (5FO-36bT
5MCCC 1A6451T) and B. pumilus DSMZ 27T (5ATCC
7061T 5 MCCC 1A06453T) originated from the Deutsche
Sammlung von Mikroorganismen und Zellkulturen (DSMZ)
and were accessioned into the Marine Culture Collection of
China (MCCC). The type strain B. xiamenensis MCCC
1A00008T (5HYC-10T) was obtained from the MCCC.
The resequenced 16S rRNA and gyrB genes of strain
Downloaded from www.microbiologyresearch.org by
IP: 88.99.165.207
On: Sun, 18 Jun 2017 23:31:58
Printed in Great Britain
2769
Y. Liu and others
B. invictae DSMZ 26896T were identical with previous results
(Branquinho et al., 2014a), suggesting reliability of this strain
in this study.
In this study, the draft genome sequence of strain
B. invictae Bi.FFUP1T was determined by Shanghai Majorbio
Bio-pharm Technology (Shanghai, China), using Solexa
paired-end (500 bp library) sequencing technology. A
total of 500 Mbp clean data were generated to reach
about 100-fold depth of coverage with an Illumina/Solexa
Genome Analyser IIx. The clean data were assembled by
SOAPdenovo2 (Luo et al., 2012). Subsequently the digital
DNA–DNA hybridization (dDDH) and average nucleotide
identity (ANI) values between strain B. invictae DSMZ
26896T and the other four type strains, B. altitudinis
41KF2bT, B. xiamenensis HYC-10T, B. safensis FO-36bT
and B. pumilus ATCC 7061T, were respectively calculated
using a web tool of the genome-to-genome distance calculator (GGDC2.0) (Meier-Kolthoff et al., 2013) and an
EzGenome web service based on the algorithm of Goris
et al. (2007). Meanwhile, similarities of gyrB gene sequence
between strain B. invictae DSMZ 26896T and the other
four type strains were determined using DNAMAN version
8 (Lynnon). In addition, phenotypic comparisons were
carried out with API 20NE, API ZYM, API 50CHB
(bioMérieux) and Biolog GN2 systems according to the
manufacturers’ instructions.
The 70 % DDH value has served as a gold standard for
species delineation in the classification of prokaryotes
(Wayne et al., 1987). Consequently, the draft genome
sequence of strain B. invictae DSMZ 26896T was obtained
(GenBank accession number: JXAI00000000). As shown
in Table 1, the dDDH values between strain B. invictae
DSMZ 26896T and B. altitudinis 41KF2bT, B. xiamenensis
HYC-10T, B. safensis FO-36bT and B. pumilus ATCC
7061T were, respectively, 82.9 %, 43.8 %, 36.5 % and
36.2 %, suggesting that strain B. invictae DSMZ 26896T
should belong to the species B. altitudinis. However, the
dDDH value between strain B. invictae DSMZ 26896T
and these four type strains were, respectively, 61.9 %,
61.7 %, 49.9 % and 39.6 % in the original study (Branquinho et al., 2014a). Generally, such discrepancy may be
caused by use of the different methods. In contrast to the
dDDH, wet-lab DDH is much more likely to yield imprecise and error-prone results owing to dependency on the
reannealing temperature used or the particular method
employed by a laboratory (Gevers et al., 2005). Therefore,
genome sequence-based dDDH has been proposed to
replace wet-lab DDH to be used in microbial taxonomy
(Auch et al., 2010; Meier-Kolthoff et al., 2013). Meanwhile,
the ANI values between strain B. invictae DSMZ 26896T
and B. altitudinis 41KF2bT, B. xiamenensis HYC-10T,
B. safensis FO-36bT and B. pumilus ATCC 7061T were
98.1 %, 91.2 %, 88.9 % and 88.6 %, respectively. According to the standard ANI criteria for species definition
(95–96 %) (Richter & Rosselló-Móra, 2009), two type
strains, B. invictae DSMZ 26896T and B. altitudinis
41KF2bT, should be attributed to the same species.
In addition, the similarity of gyrB gene sequences between
strains B. invictae DSMZ 26896T and B. altitudinis 41KF2bT
was 98.7 %, while similarities of the gyrB gene sequence
between strain B. invictae DSMZ 26896T and the other
three type strains were below 95 %. As a result, in the
light of the proposals of 95 % gyrB gene sequence similarity
corresponding to 70 % DNA–DNA relatedness among
species of the Bacillus subtilis group (Wang et al., 2007)
and an interspecies gap of 95–96 % gyrB gene sequence
similarity within the B. pumilus group (Liu et al., 2013),
strain B. invictae DSMZ 26896T fell into the species
B. altitudinis. The genomic DNA G+C content of strain
B. invictae DSMZ 26896T was 41.1 mol% based on its
draft genome sequence and accordant with the previously
reported value of 41 mol% (Branquinho et al., 2014a).
In comparison with genotype, phenotype has played a vital
role in identification and classification of bacteria and
archaea (Kämpfer, 2014). Therefore, phenotypic characteristics of all type strains within the B. pumilus group were
analysed. The results are shown in Tables 2 and 3. In API
20NE and API ZYM tests, the results of our study are
almost consistent with those of Branquinho et al. (2014a).
Two type strains, B. invictae DSMZ 26896T and B. altitudinis
41KF2bT, revealed more consistent phenotypic features relative to the other three type strains. In Biolog tests, similar
cases were also encountered. In contrast, more differences
between our results and those of Branquinho et al.
(2014a) were exhibited in API 50CHB tests. To our knowledge, inconsistent results from the same strains used in
different studies are normal and understandable. We
believed that some reasons may explain these discrepancies.
On the one hand, several results of API 50CHB in the study
Table 1. dDDH and ANI values, and gyrB gene sequence similarities between strain B. invictae DSMZ 26896T and reference
strains
Strain
B.
B.
B.
B.
B.
T
invictae DSMZ 26896
altitudinis DSMZ 21631T
pumilus ATCC 7061T
xiamenensis HYC-10T
safensis FO-36bT
2770
Accession no.
dDDH
ANIm
gyrB
JXAI00000000
ASJC00000000
ABRX01000000
AMSH00000000
ASJD00000000
100 %
82.9 %
36.2 %
43.8 %
36.5 %
100 %
98.1 %
88.6 %
91.2 %
88.9 %
100 %
98.7 %
94.3 %
91.7 %
90.8 %
Downloaded from www.microbiologyresearch.org by
International Journal of Systematic and Evolutionary Microbiology 65
IP: 88.99.165.207
On: Sun, 18 Jun 2017 23:31:58
Bacillus invictae is a later synonym of B. altitudinis
Table 2. Characteristics of the five type strains within the B. pumilus group
Strains: 1, B. invictae DSMZ 26896T; 2, B. altitudinis DSMZ 21631T; 3, B. xiamenensis HYC-10T; 4, B. safensis FO-36bT; 5, B. pumilus ATCC 7061T.
Data for strains 1 and 2 were from this study, while data for strains 3, 4 and 5 were from the study by Lai et al. (2014). In API 20NE tests, all strains
were positive for b-glucosidase (aesculin hydrolysis), gelatin hydrolysis, b-galactosidase, and utilization of D -glucose, D -mannitol, D -mannose,
malic acid, N-acetylglucosamine, potassium gluconate and trisodium citrate; all strains were negative for reduction of nitrate, denitrification,
indole production, arginine dihydrolase, urease, and utilization of adipic acid and capric acid. In API ZYM tests, all strains were positive for alkaline
phosphatase, acid phosphatase, esterase (C4), esterase lipase (C8), naphthol-AS-BI-phosphoamidase and b-glucosidase; all strains were negative for
N-acetyl-b-glucosaminidase, a-fucosidase, a-galactosidase and b-glucuronidase. In Biolog tests, all strains were able to utilize acetic acid, itaconic
acid, N-acetyl-D -glucosamine, adonitol, cellobiose, L -fucose, gentiobiose, lactulose, D -mannitol, L -rhamnose, D -sorbitol, formic acid, D -glucosaminic acid, c-hydroxybutyric acid, D -alanine, c-aminobutyric acid, inosine, uridine, thymidine and phenylethylamine; none of the strains could utilize
a-cyclodextrin, D -fructose, dextrin, D -mannose, Tween 80, N-acetyl-D -galactosamine, a-D -glucose, melibiose, methyl b-D -glucoside, sucrose, turanose, xylitol, D -galacturonic acid, D -glucuronic acid, b-hydroxybutyric acid, a-ketoglutaric acid, a-ketovaleric acid, malonic acid, D -saccharic acid,
bromosuccinic acid, succinamic acid, glucuronamide, L -alaninamide, L -glutamic acid, glycyl-L -aspartic acid, L -histidine, L -threonine, glycerol, DL a-glycerol phosphate or glucose 1-phosphate. +, Positive; 2, negative; W , weakly positive.
Characteristic
API 20NE
D -Glucose fermentation, maltose
L -Arabinose
API ZYM
Cystine aminopeptidase, trypsin
a-Mannosidase
Leucine aminopeptidase
Lipase (C14)
Valine aminopeptidase
a-Chymotrypsin
a-Glucosidase
b-Galactosidase
Biolog
cis-Aconitic acid, putrescine
i-Erythritol, L -alanine, hydroxy-L -proline
D -Psicose, citric acid
Monomethyl succinate
a-Lactose
Quinic acid
Succinic acid, D -serine
Glycogen, L -arabinose
D -Galactonic acid lactone
a-Hydroxybutyric acid
Urocanic acid
D -Arabitol
Methyl pyruvate
D,L -Lactic acid
Tween 40
Maltose
L -Alanyl glycine
D -Trehalose, a-ketobutyric acid, L -pyroglutamic acid
D -Gluconic acid, L -leucine, 2-aminoethanol
Propionic acid
L -Proline
p-Hydroxyphenylacetic acid
D -Raffinose, glucose 6-phosphate
L -Phenylalanine
L -Aspartic acid
L -Serine
D -Galactose
Sebacic acid, L -asparagine
myo-inositol
http://ijs.sgmjournals.org
1
2
3
4
5
2
+
2
+
2
2
+
+
2
+
2
2
2
2
2
W
W
+
2
+
W
W
W
W
W
W
W
W
+
+
+
+
+
+
+
W
W
W
W
W
W
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
+
+
+
+
+
+
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
2
+
+
+
2
2
2
2
2
2
2
+
+
+
+
+
+
+
W
W
2
+
W
W
W
W
W
Downloaded from www.microbiologyresearch.org by
IP: 88.99.165.207
On: Sun, 18 Jun 2017 23:31:58
W
W
W
W
W
W
2
W
2
+
W
+
+
+
2
2
W
W
2
+
2
+
2
+
+
+
W
W
W
+
+
+
+
W
+
+
+
2
W
2
W
+
+
2
W
+
W
W
W
+
W
W
2
2
2
+
+
2
+
+
+
+
+
+
+
2
+
W
2
+
2
+
2
+
+
W
+
+
+
+
2
W
2771
Y. Liu and others
Table 2. cont.
Characteristic
1
2
3
4
5
2,3-Butanediol
Glycyl-L -glutamic acid
L -Ornithine
D,L -Carnitine
DNA G+C content (mol%)
W
W
W
2
2
+
+
W
W
W
W
W
W
+
2
+
41.7
W
W
W
W
41.4
41.3
41.3
41.6
Nevertheless, in comparison with the results in Table 3, we
discovered that our results agreed almost entirely with
those from the literature, while the results of Branquinho
et al. (2014a) did not. Therefore, our results of API 50 CHB
tests of these strains are more reliable. In brief, the two strains,
B. invictae DSMZ 26896T and B. altitudinis 41KF2bT, are difficult to distinguish in terms of a small number of differences
of Branquinho et al. (2014a) are ambiguous, such as D xylose (+/2), D -galactose (+/2), methyl a-D -mannoside
(+/2) and amygdalin (+/2). On the other hand, some
strains being clones can lose some traits as a result of
mutations during their storage and growth. In addition, it
is possible to bring about some differences during manual
operation in tests and determination of results.
Table 3. Profiles of acid production from carbohydrates of the five type strains within the B. pumilus group
Strains: 1, B. invictae DSMZ 26896T; 2, B. altitudinis DSMZ 21631T; 3, B. xiamenensis MCCC 1A00008T; 4, B. safensis DSMZ 19292T; 5, B. pumilus
DSMZ 27T. Data from this study except where indicated otherwise. All strains can produce acid by use of glycerol, D -ribose, D -fructose, arbutin,
aesculin, salicin, sucrose and trehalose; none of the strains produce acid using L -xylose, D -adonitol, methyl b-D -xyloside, L -sorbose, dulcitol, inulin,
melezitose, glycogen, xylitol, D -lyxose, D -fucose, L -fucose, D -arabitol, L -arabitol, gluconate, 2-ketogluconate or 5-ketogluconate. +, Positive; 2,
negative; W , weakly positive, V , variable.
Carbohydrate
D -Turanose
Inositol
Lactose
Melibiose
Erythritol
Raffinose
Methyl-D -glucopyranoside
Maltose
L -Rhamnose
D -Sorbitol
D -Arabinose
Starch
D -Mannitol
N-Acetylglucosamine
D -Glucose
D -Tagatose
L -Arabinose
D -Mannose
Cellobiose
D -Ribose
Gentiobiose
D -Xylose
Methyl a-D -mannoside
D -Galactose
Amygdalin
1*
1
2*
2
3*
3
4*
4
4D
5*
5
5d
2
2
2
2
2
2
2
2
2
2
2
2
2
2
+
+
+
+
+
+
+
+/2
+/2
+/2
+/2
2
2
2
2
2
2
2
2
2
2
2
2
+
2
2
2
2
2
2
2
+
+
+
+
+
+
2
2
+
+
+
+
+
+
+
+
+
+
2
2
2
2
2
2
2
2
2
2
2
+
2
2
+
+
2
+
+
2
2
+
+
+
+
+
+
+
+
+
+
2
2
2
2
2
2
2
2
2
2
2
2
+
+
+
2
2
+
+
+
+
2
2
2
+
2
2
2
2
+
2
2
2
2
2
+
2
+
2
+
+
+
2
+
+
+
+
+
+
+
+
+
2
2
2
+
+
+
2
2
2
2
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
2
+
2
2
+
+
2
+
2
2
+
2
+
2
2
+
+
+
2
+
+
+
+
+
+
+
+
2
2
2
31 (+)
11 (2)
14 (2)
15 (2)
1 (2)
15 (2)
27 (+)
35 (+)
14 (2)
2 (2)
1 (2)
1 (2)
99 (+)
64 (+)
99 (+)
90 (+)
88 (+)
99 (+)
99 (+)
97 (+)
67 (+)
65 (+)
37 (+)
49 (+)
62 (+)
W
+
+
+
+
+
+
+
2
2
+
+
W
2
2
2
+
+
+
+
+
+
+
+
+
+
+
W
W
+
+
V
V
2
2
+
+
2
2
2
2
+
+
+
+
+
+
+
+
V
+
+
+
+
W
+
+
2
2
2
2
+
+
+
+
+
+
+
+
+
+
W
+
+
*Data from earlier results by Branquinho et al. (2014a).
DData from earlier results by Satomi et al. (2006).
dData from http://www.biologiemarine.com/___fiches/APIpdf/api50%20CHBE%20Medium-_07964_-_F_-_50430.pdf (page 44).
2772
Downloaded from www.microbiologyresearch.org by
International Journal of Systematic and Evolutionary Microbiology 65
IP: 88.99.165.207
On: Sun, 18 Jun 2017 23:31:58
Bacillus invictae is a later synonym of B. altitudinis
in phenotypic characteristics relative to the other three type
strains within the B. pumilus group (Tables 2 and 3).
On the basis of the genotypic and phenotypic evidence
described above, we propose that B. invictae (Branquinho
et al., 2014a) should be regarded as a later heterotypic
synonym of B. altitudinis (Shivaji et al., 2006). According
to the rules of priority (Rules 38 and 42 of the Bacteriological Code; Lapage et al., 1992), the name Bacillus altitudinis
is retained for the united taxon, with Bacillus invictae as a
later heterotypic synonym.
values and their relationship to whole-genome sequence similarities.
Int J Syst Evol Microbiol 57, 81–91.
Kämpfer, P. (2014). Continuing importance of the ‘‘phenotype’’ in
the genomic era. Methods Microbiol 41, 307–320.
Lai, Q., Liu, Y. & Shao, Z. (2014). Bacillus xiamenensis sp. nov., isolated
from intestinal tract contents of a flathead mullet (Mugil cephalus).
Antonie van Leeuwenhoek 105, 99–107.
Lapage, S. P., Sneath, P. H. A., Lessel, E. F., Skerman, V. B. D.,
Seeliger, H. P. R. & Clark, W. A. (1992). International Code of
Nomenclature of Bacteria (1990 Revision). Bacteriological Code.
Washington, DC: American Society for Microbiology.
Liu, Y., Lai, Q., Dong, C., Sun, F., Wang, L., Li, G. & Shao, Z. (2013).
Phylogenetic diversity of the Bacillus pumilus group and the marine
ecotype revealed by multilocus sequence analysis. PLoS One 8, e80097.
Acknowledgements
This work was supported financially by the National Infrastructure of
Microbial Resources of China (nos NIMR-2014-9; NIMR-2014-9).
Luo, R., Liu, B., Xie, Y., Li, Z., Huang, W., Yuan, J., He, G., Chen, Y.,
Pan, Q. & other authors (2012). SOAPdenovo2: an empirically
improved memory-efficient short-read de novo assembler. Gigascience
1, 18.
Meier-Kolthoff, J. P., Auch, A. F., Klenk, H. P. & Göker, M. (2013).
References
Auch, A. F., von Jan, M., Klenk, H. P. & Göker, M. (2010). Digital
Genome sequence-based species delimitation with confidence intervals and improved distance functions. BMC Bioinformatics 14, 60.
DNA-DNA hybridization for microbial species delineation by means
of genome-to-genome sequence comparison. Stand Genomic Sci 2,
117–134.
Richter, M. & Rosselló-Móra, R. (2009). Shifting the genomic gold
Branquinho, R., Sousa, C., Osório, H., Meirinhos-Soares, L., Lopes,
J., Carriço, J. A., Busse, H. J., Abdulmawjood, A., Klein, G. & other
authors (2014a). Bacillus invictae sp. nov., isolated from a health
Satomi, M., La Duc, M. T. & Venkateswaran, K. (2006). Bacillus
product. Int J Syst Evol Microbiol 64, 3867–3876.
Shivaji, S., Chaturvedi, P., Suresh, K., Reddy, G. S., Dutt, C. B.,
Wainwright, M., Narlikar, J. V. & Bhargava, P. M. (2006). Bacillus
Branquinho, R., Meirinhos-Soares, L., Carriço, J. A., Pintado, M. &
Peixe, L. V. (2014b). Phylogenetic and clonality analysis of Bacillus
standard for the prokaryotic species definition. Proc Natl Acad Sci
U S A 106, 19126–19131.
safensis sp. nov., isolated from spacecraft and assembly-facility
surfaces. Int J Syst Evol Microbiol 56, 1735–1740.
pumilus isolates uncovered a highly heterogeneous population of
different closely related species and clones. FEMS Microbiol Ecol 90,
689–698.
aerius sp. nov., Bacillus aerophilus sp. nov., Bacillus stratosphericus
sp. nov. and Bacillus altitudinis sp. nov., isolated from cryogenic
tubes used for collecting air samples from high altitudes. Int J Syst
Evol Microbiol 56, 1465–1473.
Branquinho, R., Sousa, C., Lopes, J., Pintado, M. E., Peixe, L. V. &
Osório, H. (2014c). Differentiation of Bacillus pumilus and Bacillus
Wang, L. T., Lee, F. L., Tai, C. J. & Kasai, H. (2007). Comparison of
safensis using MALDI-TOF-MS. PLoS One 9, e110127.
Gevers, D., Cohan, F. M., Lawrence, J. G., Spratt, B. G., Coenye, T.,
Feil, E. J., Stackebrandt, E., Van de Peer, Y., Vandamme, P. &
other authors (2005). Opinion: Re-evaluating prokaryotic species.
gyrB gene sequences, 16S rRNA gene sequences and DNA-DNA
hybridization in the Bacillus subtilis group. Int J Syst Evol Microbiol
57, 1846–1850.
Nat Rev Microbiol 3, 733–739.
Wayne, L., Brenner, D., Colwell, R., Grimont, P., Kandler, O.,
Krichevsky, M., Moore, L., Moore, W., Murray, R. & other authors
(1987). International Committee on Systematic Bacteriology. Report
Goris, J., Konstantinidis, K. T., Klappenbach, J. A., Coenye, T.,
Vandamme, P. & Tiedje, J. M. (2007). DNA-DNA hybridization
of the ad hoc committee on reconciliation of approaches to
bacterial systematics. Int J Syst Bacteriol 37, 463–464.
http://ijs.sgmjournals.org
Downloaded from www.microbiologyresearch.org by
IP: 88.99.165.207
On: Sun, 18 Jun 2017 23:31:58
2773