International Journal of Systematic and Evolutionary Microbiology (2013), 63, 3704–3709
DOI 10.1099/ijs.0.049734-0
Algibacter undariae sp. nov., isolated from a brown
algae reservoir
Sooyeon Park,1 Jung-Sook Lee,2 Keun-chul Lee2 and Jung-Hoon Yoon1
Correspondence
J.-H. Yoon
[email protected]
1
Department of Food Science and Biotechnology, Sungkyunkwan University, Jangan-gu, Suwon,
South Korea
2
Korea Research Institute of Bioscience and Biotechnology (KRIBB), PO Box 115, Yuseong,
Daejeon, South Korea
A Gram-stain-negative, non-flagellated, rod-shaped bacterial strain able to move by gliding,
designated WS-MY9T, was isolated from a brown algae reservoir in South Korea. Strain WSMY9T grew optimally at 25 6C, at pH 7.0–8.0 and in the presence of 2 % (w/v) NaCl. A
neighbour-joining phylogenetic tree based on 16S rRNA gene sequences showed that strain
WS-MY9T clustered with the type strain of Algibacter lectus with a bootstrap resampling value of
100 %. Strain WS-MY9T exhibited 16S rRNA gene sequence similarity values of 98.5 and 96.7 %
to the type strains of A. lectus and Algibacter mikhailovii, respectively, and less than 96.1 %
sequence similarity to other members of the family Flavobacteriaceae. Strain WS-MY9T contained
MK-6 as the predominant menaquinone and anteiso-C15 : 0, iso-C17 : 0 3-OH, iso-C15 : 1 G and
iso-C15 : 0 as the major fatty acids. The major polar lipids of strain WS-MY9T were
phosphatidylethanolamine and two unidentified lipids. The DNA G+C content of strain WSMY9T was 35.0 mol% and its DNA–DNA relatedness value with A. lectus KCTC 12103T was
15 %. The phylogenetic and genetic distinctiveness and differential phenotypic properties
revealed that strain WS-MY9T is separate from the two recognized species of the genus
Algibacter. On the basis of the data presented, strain WS-MY9T represents a novel species of the
genus Algibacter, for which the name Algibacter undariae sp. nov. is proposed. The type strain is
WS-MY9T (5KCTC 32259T5CCUG 63684T).
The genus Algibacter, a member of the family Flavobacteriaceae in the phylum Bacteroidetes (Bernardet, 2011),
was first proposed by Nedashkovskaya et al. (2004) with
Algibacter lectus, isolated from green algae, as the type
species. The second species of the genus Algibacter,
Algibacter mikhailovii, was subsequently described following isolation from a sea urchin (Nedashkovskaya et al.,
2007). Recently, Algibacter aquimarinus has been described
and Algibacter pectinivorans reclassified as a member of the
genus Algibacter (Park et al., 2013). In this study, a novel
bacterial strain, designated WS-MY9T and isolated from
brown algae collected from a reservoir located on Wando,
an island of South Korea, is described. Comparative 16S
rRNA gene sequence analysis indicated that strain WSMY9T was most closely related to members of the genus
Algibacter. The aim of the present work was to determine
the exact taxonomic position of strain WS-MY9T by using
a polyphasic characterization that included the determination of phenotypic properties, a detailed phylogenetic
The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene
sequence of strain WS-MY9T is KC261664.
A supplementary figure is available with the online version of this paper.
3704
investigation based on 16S rRNA gene sequences and
genetic analysis.
Strain WS-MY9T was isolated by the dilution plating
technique at 25 uC on marine agar 2216 (MA; BD Difco)
and cultivated routinely under the same conditions.
Algibacter lectus KCTC 12103T and Algibacter mikhailovii
LMG 23988T were used as reference strains for DNA–DNA
hybridization, fatty acid and polar lipid analyses and a
number of other phenotypic characterizations. Cell morphology and flagellation were examined by using light
microscopy (BX51; Olympus) and transmission electron
microscopy (JEM1010; JEOL), respectively. To assess
flagellation, cells from an exponentially growing MA
culture were negatively stained with 1 % (w/v) phosphotungstic acid and the grids were examined after being airdried. Gliding motility was investigated as described by
Bowman (2000). The Gram reaction was determined by
using the bioMérieux Gram-stain kit according to the
manufacturer’s instructions. Growth under anaerobic
conditions was determined after incubation for 10 days
in an anaerobic jar (MGC) with AnaeroPack (MGC) on
MA; the jar was kept overnight at 4 uC to create anoxic
conditions before incubation at 25 uC. Growth at 4, 10, 20,
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Algibacter undariae sp. nov.
25, 28, 30, 35, 37 and 40 uC was measured on MA. The pH
range for growth was investigated in marine broth 2216
(MB; BD Difco) adjusted to pH 4.5–9.5 (in increments of
0.5 pH unit) by using sodium acetate/acetic acid and
sodium carbonate buffers. The pH values were verified
after autoclaving. Growth in the absence of NaCl and in the
presence of 0.5, 1.0, 2.0 and 3.0 % (w/v) NaCl was
investigated in trypticase soy broth prepared according to
the formula of the BD Bacto medium except that NaCl was
excluded and that 0.45 % (w/v) MgCl2 . 6H2O was added.
Growth in the presence of 2.0–8.0 % (as final concentration, w/v, in increments of 1.0 %) NaCl was investigated in
MB. Catalase and oxidase activities were determined as
described by Lányı́ (1987). Hydrolysis of casein, starch,
hypoxanthine, L-tyrosine and xanthine was tested on MA
using the substrate concentrations described by Barrow &
Feltham (1993). Hydrolysis of gelatin and urea was
investigated by using Nutrient Gelatin medium (BD
Difco) and Urea Agar Base medium (BD BBL), respectively, with the modification that artificial seawater was used
for the preparation of media. Hydrolysis of aesculin and
Tweens 20, 40, 60 and 80, and nitrate reduction were
investigated as described by Lányı́ (1987) with the modification that artificial seawater was used for preparation of
media. The artificial seawater contained (l21 distilled
water) 23.6 g NaCl, 0.64 g KCl, 4.53 g MgCl2 . 6H2O,
5.94 g MgSO4 . 7H2O and 1.3 g CaCl2 . 2H2O (Bruns et al.,
2001). H2S production was tested as described by Bruns
et al. (2001). The presence of flexirubin-type pigments was
investigated as described by Reichenbach (1992) and
Bernardet et al. (2002). Utilization of substrates as sole
carbon and energy sources was tested as described by
Baumann & Baumann (1981), using supplementation with
2 % (v/v) Hutner’s mineral base (Cohen-Bazire et al., 1957)
and 1 % (v/v) vitamin solution (Staley, 1968). Acid
production from carbohydrates was tested as described
by Leifson (1963). Susceptibility to antibiotics was tested
on MA plates using antibiotic discs (Advantec) containing
(mg per disc unless otherwise stated): ampicillin (10),
carbenicillin (100), cephalothin (30), chloramphenicol
(100), gentamicin (30), kanamycin (30), lincomycin (15),
neomycin (30), novobiocin (5), oleandomycin (15), penicillin G (20 U), polymyxin B (100 U), streptomycin (50)
and tetracycline (30). Enzyme activities were determined
using the API ZYM system (bioMérieux), with the test strips
incubated for 8 h at 25 uC. Morphological, cultural,
physiological and biochemical properties of strain WSMY9T are given in the species description and in Table 1.
Cell biomass of strain WS-MY9T for DNA extraction and
for the analyses of isoprenoid quinones and polar lipids
was obtained from cultures grown at 25 uC in MB. Cell
biomass of A. lectus KCTC 12103T and A. mikhailovii LMG
23988T for DNA extraction and for polar lipid analysis
was obtained from cultures grown at 25 uC in MB.
Chromosomal DNA was isolated and purified according
to the method described by Yoon et al. (1996), with the
exception that RNase T1 was used in combination with
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RNase A to minimize contamination by RNA. The 16S
rRNA gene was amplified by PCR as described previously
(Yoon et al., 1998) using two universal primers, 9F (59GAGTTTGATCCTGGCTCAG-39) and 1512R (59-ACGGTTACCTTGTTACGACTT-39), and the PCR products
were purified by using a QIAquick PCR purification kit
(Qiagen). Sequencing of the amplified 16S rRNA gene and
phylogenetic analysis were performed as described by
Yoon et al. (2003). Isoprenoid quinones were extracted
according to the method of Komagata & Suzuki (1987)
and analysed using reversed-phase HPLC and a YMC
ODS-A (25064.6 mm) column. The isoprenoid quinones
were eluted by a mixture of methanol/2-propanol (2 : 1,
v/v), using a flow rate of 1 ml min21 at room temperature
and detected by UV absorbance at 270 nm. For cellular
fatty acid analysis, cell mass of strain WS-MY9T, A. lectus
KCTC 12103T and A. mikhailovii LMG 23988T was
harvested from MA plates after cultivation for 3 days at
25 uC. The physiological age of the cell masses was
standardized by observing the growth development of cells
on the agar plates followed by harvesting them from the
same quadrant on the plates for each strain according
to the standard MIDI protocol (Sherlock Microbial
Identification System, version 6.1). Fatty acids were
saponified, methylated and extracted using the standard
MIDI protocol (Sherlock Microbial Identification System,
version 6.1). The fatty acids were analysed by GC (6890;
Hewlett Packard) and identified by using the TSBA6
database of the Microbial Identification System (Sasser,
1990). Polar lipids were extracted according to the
procedures described by Minnikin et al. (1984), and
separated by two-dimensional TLC using chloroform/
methanol/water (65 : 25 : 3.8, by vol.) for the first dimension and chloroform/methanol/acetic acid/water (40 : 7.5 :
6 : 1.8, by vol.) for the second dimension as described by
Minnikin et al. (1977). Individual polar lipids were
identified by spraying the TLC plates with molybdophosphoric acid, molybdenum blue, ninhydrin and anaphthol reagents (Minnikin et al., 1984; Komagata &
Suzuki, 1987) and with Dragendorff’s reagent (Sigma).
The DNA G+C content was determined by the method of
Tamaoka & Komagata (1984) with the modification that
DNA was hydrolysed and the resultant nucleotides were
analysed by reversed-phase HPLC equipped with a YMC
ODS-A (25064.6 mm) column. The nucleotides were
eluted by a mixture of 0.55 M NH4H2PO4 (pH 4.0) and
acetonitrile (40 : 1, v/v), using a flow rate of 1 ml min21 at
room temperature and detected by UV absorbance at
270 nm. DNA–DNA hybridization was performed fluorometrically at 45 uC by the method of Ezaki et al. (1989)
using photobiotin-labelled DNA probes and microdilution wells. Hybridization was performed with five
replications for each sample. DNA of strain WS-MY9T
and A. lectus KCTC 12103T was used as probes for
reciprocal hybridizations. The highest and lowest values
obtained for each sample were excluded and the means of
the remaining three values were quoted as DNA–DNA
relatedness values.
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S. Park and others
Table 1. Differential phenotypic characteristics of strain WSMY9T and the type strains of two species of the genus
Algibacter
Strains: 1, WS-MY9T; 2, A. lectus KCTC 12103T; 3, A. mikhailovii
LMG 23988T. Data obtained from this study unless otherwise
indicated. All strains are positive for the following: gliding motility*;
catalase and oxidase activities*; hydrolysis of L-tyrosine; utilization of
D-galactose, D-glucose, D-mannose and D-xylose; acid production
from D-galactose, D-glucose, lactose, D-mannose and D-xylose;
susceptibility to chloramphenicol, lincomycin, novobiocin and
oleandomycin; and activity of alkaline phosphatase, [esterase (C4)
and esterase lipase (C8) (weak for A. mikhailovii LMG 23988T)],
leucine arylamidase and acid phosphatase. All strains are negative for
the following: Gram stain; production of flexirubin-type pigments*;
H2S production; hydrolysis of casein*, hypoxanthine, urea and
xanthine; utilization of L-arabinose, trehalose, acetate, benzoate,
citrate, formate, L-malate, pyruvate, succinate, L-glutamate and
salicin; acid production from L-arabinose, melezitose, D-ribose,
trehalose, myo-inositol and D-sorbitol; susceptibility to ampicillin,
gentamicin, kanamycin, neomycin, polymyxin B and streptomycin;
and activity of lipase (C14), trypsin, a-chymotrypsin, a-galactosidase,
b-glucuronidase, b-glucosidase, a-mannosidase and a-fucosidase. +,
Positive; –, negative; w, weakly positive.
Characteristic
1
2
3
Growth at 37 uC
Anaerobic growth
Nitrate reduction
Hydrolysis of:
Aesculin
Gelatin
Starch
Tweens 20 and 40
Tween 60
Tween 80
Utilization of:
Cellobiose
D-Fructose
Maltose
Sucrose
Acid production from:
Cellobiose
D-Fructose
Maltose
Melibiose
Raffinose
L-Rhamnose
Sucrose
D-Mannitol
Susceptibility to:
Carbenicillin
Cephalothin
Penicillin G
Tetracycline
Enzyme activity (API ZYM)
Valine arylamidase
Cystine arylamidase
–
–
–
–*
+*
–*
+*
–*
+*
–
–
–
+
+
+
+
+*
+*
+*
+
–*
+
+*
–*
–*
–
–*
–
+
–
–
+
+
+
+
+
–
+
–
–
+
–
+
–
–
–
+
+
+
+
+
+
+
+
+
+
–
+
–
–
–
–
–
+
+
+
–
+
+
+
+
–
–
–
+
+
–
–
–
+
w
3706
Table 1. cont.
Characteristic
Naphthol-AS-BI-phosphohydrolase
b-Galactosidase
a-Glucosidase
N-Acetyl-b-glucosaminidase
DNA G+C content (mol%)
1
2
3
+
–
+
+
35.0
+
+
–
+
31.9*
–
–
–
–
35.1*
*Data for reference strains taken from Nedashkovskaya et al. (2004,
2007).
The almost-complete 16S rRNA gene sequence of strain
WS-MY9T comprised 1441 nt, representing approximately
95 % of the Escherichia coli 16S rRNA gene sequence. In the
phylogenetic tree reconstructed using the neighbour-joining algorithm, strain WS-MY9T clustered with A. lectus
KMM 3902T with a bootstrap resampling value of 100 %,
and this cluster joined the cluster comprising the type
strains of A. mikhailovii, Algibacter aquimarinus, Algibacter
pectinivorans and Postechiella marina (Fig. 1). The
relationship between strain WS-MY9T and A. lectus
KMM 3902T was also maintained in the trees reconstructed
using the maximum-likelihood and maximum-parsimony
algorithms (Fig. 1). However, the cluster comprising strain
WS-MY9T and A. lectus KMM 3902T might not form a
monophyletic group with the cluster comprising the type
strains of A. mikhailovii, A. aquimarinus and Algibacter
pectinivorans, depending on which species are selected for
the phylogenetic analysis (data not shown). Strain WSMY9T exhibited 16S rRNA gene sequence similarity values
of 98.5, 96.7, 94.8, 96.1 and 94.2 % to A. lectus KMM
3902T, A. mikhailovii KMM 6171T, A. aquimarinus
KYW589T, Algibacter pectinivorans JC2675T and
Postechiella marina M091T, respectively, and of less than
95.5 % to those of the other species.
The predominant isoprenoid quinone detected in strain
WS-MY9T was menaquinone-6 (MK-6) in line with the
genus Algibacter (Nedashkovskaya et al., 2004) and all
other members of the family Flavobacteriaceae (Bernardet,
2011). The complete cellular fatty acid profiles of strain
WS-MY9T and the type strains of the two species of the
genus Algibacter with validly published names at the time
of this study (A. lectus KCTC 12103T and A. mikhailovii
LMG 23988T) are compared in Table 2. The major fatty
acids (.10 % of the total fatty acids) found in strain WSMY9T were anteiso-C15 : 0 (19.8 %), iso-C17 : 0 3-OH
(15.6 %), iso-C15 : 1 G (12.4 %) and iso-C15 : 0 (11.3 %)
(Table 2). The fatty acid profiles of the three strains grown
and analysed under identical conditions in this study were
similar, but there were also differences in the proportions
of some fatty acids. A. mikhailovii LMG 23988T was
characterized by having a higher amount of summed
feature 3 (C16 : 1v7c and/or C16 : 1v6c), which was also
shown in the study of Park et al. (2013). Strain WS-MY9T
contained phosphatidylethanolamine (PE) and two
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Algibacter undariae sp. nov.
Algibacter aquimarinus KYW589T (JQ259859)
50.5
50.6
Algibacter mikhailovii KMM 6171T (AM491809)
0.01
Algibacter pectinivorans JC2675T (HM475134)
Postechiella marina M091T (HQ336487)
Algibacter undariae WS-MY9T (KC261664)
100
Algibacter lectus KMM 3902T (AY187689)
Tamlana agarivorans JW-26T (EU221275)
96.3
79.0
Tamlana crocina HST1-43T (AM286230)
Gaetbulibacter saemankumensis SMK-12T (AY883937)
Gaetbulibacter marinus IMCC1914T (EF108219)
Flaviramulus basaltis H35T (DQ361033)
Mariniflexile aquimaris HWR-17T (HQ144198)
100
Mariniflexile gromovii KMM 6038T (DQ312294)
Olleya marilimosa CAM030T (EF660466)
99.9
Olleya aquimaris L-4T (FJ886713)
Flavivirga amylovorans JC2681T (HM475138)
97.2
Flavivirga jejuensis JC2682T (HM475139)
Psychroserpens mesophilus KOPRI 13649T (DQ001321)
65.3
54.2
57.6
Psychroserpens burtonensis ACAM 188T (U62913)
100
Formosa agariphila KMM 3901T (AY187688)
Formosa algae KMM 3553T (AY228461)
Bizionia echini KMM 6177T (FJ716799)
Bizionia paragorgiae KMM 6029T (AY651070)
Capnocytophaga ochracea ATCC 27872T (U41350)
Fig. 1. Neighbour-joining tree based on 16S rRNA gene sequences showing the phylogenetic positions of strain WS-MY9T,
the type strains of species of the genus Algibacter and representatives of some other related taxa. Bootstrap values (expressed
as percentages of 1000 replications) .50 % are shown at branching points. Filled circles indicate that the corresponding
nodes were also recovered in the trees generated with the maximum-likelihood and maximum-parsimony algorithms.
Capnocytophaga ochracea ATCC 27872T (GenBank accession no. U41350) was used as an outgroup. Bar, 0.01 substitutions
per nucleotide position.
unidentified lipids (L1 and L2) as the major polar lipids; it
also contained minor amounts of two additional unidentified lipids (L3 and L4), two unidentified glycolipids and
one unidentified aminolipid (Fig. S1 available in IJSEM
Online). This polar lipid profile was similar to those of A.
lectus KCTC 12103T and A. mikhailovii LMG 23988T in
that phosphatidylethanolamine and two unidentified lipids
were major polar lipids (Fig. S1). However, one major
unidentified lipid (L2) of strain WS-MY9T and A. lectus
KCTC 12103T differed from that (L5) detected in A.
mikhailovii LMG 23988T. The DNA G+C content of strain
WS-MY9T was 35.0 mol%.
Taking these phylogenetic and chemotaxonomic data into
account, it is reasonable to classify strain WS-MY9T as a
member of the genus Algibacter. However, even the
taxonomic position of A. mikhailovii as well as the
description of Algibacter aquimarinus and reclassification
of Algibacter pectinivorans may have to be re-evaluated.
Strain WS-MY9T exhibited a mean DNA–DNA relatedness
value of 15 % with A. lectus KCTC 12103T. Strain WSMY9T was distinguishable from A. lectus KCTC 12103T and
http://ijs.sgmjournals.org
A. mikhailovii LMG 23988T by differences in several
phenotypic characteristics as shown in Table 1. These
differences, in combination with phylogenetic and genetic
distinctiveness, were sufficient to prove that the novel
strain was separate from other species of the genus
Algibacter (Wayne et al., 1987; Stackebrandt & Goebel,
1994). On the basis of these phenotypic, chemotaxonomic,
phylogenetic and genetic data, strain WS-MY9T represents
a novel species of the genus Algibacter, for which the name
Algibacter undariae sp. nov. is proposed.
Description of Algibacter undariae sp. nov.
Algibacter undariae (un.da.ri9a.e. N.L. gen. n. undariae of
Undaria, named after the generic name of the brown algae
Undaria pinnatifida, from whose reservoir the type strain
was isolated).
Cells are Gram-stain-negative, non-flagellated, motile by
gliding and rod-shaped, approximately 0.4–0.7 mm in
diameter and 0.7–6.0 mm in length. Colonies on MA are
circular, slightly convex, smooth, glistening, yellow and
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S. Park and others
Table 2. Cellular fatty acid compositions (%) of strain WSMY9T and the type strains of two species of the genus
Algibacter
Strains: 1, WS-MY9T; 2. A. lectus KCTC 12103T; 3, A. mikhailovii
LMG 23988T. All data from this study. Fatty acids that represented
,0.5 % in all strains were omitted. TR, Traces (,0.5 %); –, not
detected.
Fatty acid
Straight-chain
C16 : 0
Unsaturated
C15 : 1v6c
C17 : 1v6c
C18 : 1v5c
Branched
iso-C13 : 0
iso-C14 : 0
iso-C15 : 0
anteiso-C15 : 0
iso-C15 : 1 G*
anteiso-C15 : 1 A*
iso-C16 : 0
anteiso-C17 : 1 A*
iso-C17 : 1v9c
Hydroxy
iso-C13 : 0 3-OH
C15 : 0 2-OH
C15 : 0 3-OH
iso-C15 : 0 3-OH
C16 : 0 3-OH
iso-C16 : 0 3-OH
C17 : 0 2-OH
C17 : 0 3-OH
iso-C17 : 0 3-OH
10-Methyl C18 : 0 (TBSA)
Summed feature 3D
1
2
3
1.9
1.4
2.9
3.3
1.4
1.3
6.7
3.1
1.1
2.2
1.3
2.2
0.7
–
11.3
19.8
12.4
2.9
–
–
–
1.6
0.7
9.3
7.7
16.9
2.0
–
0.8
–
–
13.5
5.1
19.5
3.4
1.0
0.9
1.3
–
2.5
–
9.3
3.1
5.0
6.4
–
15.6
–
3.4
1.8
1.6
4.3
9.7
4.0
4.0
3.3
1.0
11.9
1.3
5.5
–
2.3
–
5.8
–
1.1
5.0
–
15.6
–
17.1
TR
citrate, formate, L-malate, pyruvate, succinate, L-glutamate
and salicin are not. Acid is produced from D-fructose,
D-galactose, D-glucose, lactose, D-mannose, melibiose, Dxylose and D-mannitol, but not from L-arabinose, cellobiose, maltose, melezitose, raffinose, L-rhamnose, D-ribose,
sucrose, trehalose, myo-inositol or D-sorbitol. Susceptible
to carbenicillin, cephalothin, chloramphenicol, lincomycin,
novobiocin, oleandomycin and penicillin G, but not to
ampicillin, gentamicin, kanamycin, neomycin, polymyxin
B, streptomycin or tetracycline. In assays with the API
ZYM system, alkaline phosphatase, esterase (C4), esterase
lipase (C8), leucine arylamidase, valine arylamidase, acid
phosphatase, naphthol-AS-BI-phosphohydrolase, a-glucosidase and N-acetyl-b-glucosaminidase activities are present, but lipase (C14), cystine arylamidase, trypsin,
a-chymotrypsin, a-galactosidase, b-galactosidase, b-glucuronidase, b-glucosidase, a-mannosidase and a-fucosidase
activities are absent. The predominant menaquinone is
MK-6. The major fatty acids (.10 % of the total fatty
acids) are anteiso-C15 : 0, iso-C17 : 0 3-OH, iso-C15 : 1 G and
iso-C15 : 0. The major polar lipids are phosphatidylethanolamine and two unidentified lipids.
The type strain, WS-MY9T (5KCTC 32259T5CCUG
63684T), was isolated from a brown algae reservoir at
Wando in the South Sea, South Korea. The DNA G+C
content of the type strain is 35.0 mol%.
Acknowledgements
This work was supported by the Program for Collection of Domestic
Biological Resources from the National Institute of Biological
Resources (NIBR) and the Program for Collection, Management
and Utilization of Biological Resources and BK 21 program from the
Ministry of Education, Science and Technology (MEST) of the
Republic of Korea.
References
*Double bond position indicated by a capital letter is unknown.
DSummed feature 3 contained C16 : 1v7c and/or C16 : 1v6c.
1.0–1.5 mm in diameter after incubation for 3 days at
25 uC. The optimal growth temperature is 25 uC; growth
occurs at 4 and 35 uC, but not at 37 uC. The optimal pH for
growth is between 7.0 and 8.0; growth occurs at pH 5.0,
but not at pH 4.5. Growth occurs in the presence of 0.5–
6.0 % (w/v) NaCl with an optimum of approximately 2.0 %
(w/v) NaCl. Mg2+ ions are required for growth. Anaerobic
growth does not occur on MA or on MA supplemented
with nitrate. H2S is not produced. Flexirubin-type
pigments are not produced. Catalase- and oxidase-positive.
Nitrate is not reduced to nitrite. Tweens 20, 40, 60 and 80
and L-tyrosine are hydrolysed, but aesculin, gelatin, casein,
hypoxanthine, starch, urea and xanthine are not. DFructose, D-galactose, D-glucose, D-mannose and D-xylose
are utilized as carbon and energy sources, but L-arabinose,
cellobiose, maltose, sucrose, trehalose, acetate, benzoate,
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