International Journal of Systematic and Evolutionary Microbiology (2014), 64, 2370–2375
DOI 10.1099/ijs.0.061747-0
Loktanella maritima sp. nov. isolated from shallow
marine sediments
Naoto Tanaka,1 Lyudmila A. Romanenko,2 Valeriya V. Kurilenko,2
Vassilii I. Svetashev,3 Natalia I. Kalinovskaya2 and Valery V. Mikhailov2
Correspondence
Naoto Tanaka
[email protected]
1
NODAI Culture Collection Center, Tokyo University of Agriculture, 1-1-1 Sakuragaoka,
Setagaya-ku, Tokyo 156-8502, Japan
2
G.B. Elyakov Pacific Institute of Bioorganic Chemistry, Far Eastern Branch, Russian Academy of
Sciences, Prospect 100 Let Vladivostoku, 159, Vladivostok 690022, Russia
3
Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences,
Vladivostok 690041, Russia
An aerobic, Gram-stain-negative, non-motile bacterium, KMM 9530T, was isolated from a
sediment sample collected from the Sea of Japan seashore. Comparative 16S rRNA gene
sequence analysis positioned novel strain KMM 9530T in the genus Loktanella as a separate line
adjacent to Loktanella sediminilitoris KCTC 32383T, Loktanella tamlensis JCM 14020T and
Loktanella maricola JCM 14564T with 98.5–98.2 % sequence similarity. Strain KMM 9530T was
characterized by its weak hydrolytic capacity and inability to assimilate most organic substrates.
The major isoprenoid quinone was Q-10, polar lipids consisted of phosphatidylcholine,
phosphatidylglycerol, diphosphatidylglycerol, an unknown phospholipid, an unknown aminolipid
and unknown lipids, and the major fatty acid was C18 : 1v7c. On the basis of phylogenetic
analysis, DNA–DNA hybridization and phenotypic characterization, it can be concluded that the
novel strain KMM 9530T represents a novel species in the genus Loktanella, for which the name
Loktanella maritima sp. nov. is proposed. The type strain of the species is KMM 9530T (5NRIC
0919T5JCM 19807T).
The genus Loktanella was initially proposed by Van
Trappen et al. (2004) to accommodate three species,
Loktanella salsilacus, as the type species of the genus,
Loktanella vestfoldensis and Loktanella fryxellensis, and
subsequently emended by Moon et al. (2010), Lee (2012)
and Tsubouchi et al. (2013). The genus Loktanella was
expanded to include twelve more species, including
Loktanella hongkongensis (Lau et al., 2004), L. agnita, L.
rosea (Ivanova et al., 2005), L. koreensis (Weon et al., 2006),
L. atrilutea (Hosoya & Yokota, 2007), L. maricola (Yoon
et al., 2007), L. pyoseonensis (Moon et al., 2010), L.
tamlensis (Lee, 2012), L. cinnabarina (Tsubouchi et al.,
2013), L. litorea (Yoon et al., 2013), L. sediminilitoris (Park
et al., 2013a) and L. soesokkakensis (Park et al., 2013b).
Here we report the polyphasic characterization of a Gramstain-negative, aerobic, beige-pigmented, non-motile bacterium, designated KMM 9530T, which was isolated from a
Abbreviations: ASW, artificial seawater; DPG, diphosphatidylglycerol;
PC, phosphatidylcholine; PG, phosphatidylglycerol.
The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene
sequence of KMM 9530T is AB894236.
Two supplementary figures are available with the online version of this
paper.
2370
shallow sediment sample. Phylogenetic analysis based on
16S rRNA gene sequence showed that strain KMM 9530T
belongs to the genus Loktanella and may represent a novel
species of this genus. Differential phenotypic properties,
together with phylogenetic distinctiveness and DNA
relatedness, demonstrated that strain KMM 9530T differed
from related species of the genus Loktanella. On the basis of
phenotypic and molecular data obtained, a novel species,
Loktanella maritima sp. nov., is described.
Strain KMM 9530T was isolated from a shallow sediment
sample, collected from Peter the Great Bay, the Sea of
Japan, Russia, as described previously (Romanenko et al.,
2004). Strain KMM 9530T was grown aerobically on/in
marine 2216 agar (MA) or marine broth (MB; BD Difco),
and stored at 280 uC in MB supplemented with 30 % (v/v)
glycerol. The type strains L. sediminilitoris KCTC 32383T, L.
maricola JCM 14564T, L. tamlensis JCM 14020T and L.
rosea KMM 6003T were kindly provided by the respective
culture collections and used in the phenotypic and lipids
analyses. Gram staining, oxidase and catalase reactions, and
motility (hanging drop method) were tested as described
by Gerhardt et al. (1994). The morphology of cells
negatively stained with 1 % phosphotungstic acid was
examined by electronic transmission microscopy [Libra 200
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Loktanella maritima sp. nov. from shallow marine sediments
FE (Carl Zeiss), provided by the Institute of Marine
Biology, Far Eastern Branch, Russian Academy of Sciences]
using carbon-coated 200-mesh copper grids. Nitrate
reduction was determined for strains grown in nitrate
broth (HiMedia) supplemented with artificial seawater
(ASW) (sulfanilic acid/a-naphthylamine test). ASW was
prepared according to the formula of Bruns et al. (2001)
using 30 g NaCl l21. Formation of H2S from thiosulfate
was tested on the ASW-based medium (2.5 g Bactopeptone
l21, 2 g yeast extract l21, 0.3 g sodium thiosulfate l21)
using a lead acetate paper strip. Hydrolysis of L-tyrosine,
chitin, xanthine and hypoxanthine was investigated by
observing transparent zones on MA supplemented with
each compound at a concentration of 1 %. Hydrolysis of
DNA was examined using DNase Test Agar (BD BBL)
prepared with ASW. Citrate utilization was tested on
Simmons’ citrate agar (HiMedia) supplemented with ASW.
Degradation of starch (0.2 %, w/v) and Tween 80 (1 %, w/
v) was tested with ASW-based basal medium, containing
5 g Bactopeptone l21, 1 g yeast extract l21, 0.1 g K2HPO4
l21 and 15 g agar l21. Hydrolysis of gelatin (0.4 %, w/v)
and casein (10 % skimmed milk, w/v; BD Difco) was
examined on ASW-based agar medium. Requirement for
and tolerance of sodium chloride was tested on ASW-based
medium using various concentrations of NaCl in the range
0–20 %, supplemented with (per litre): 10.0 g Bacto peptone,
2.0 g yeast extract, 0.028 g FeSO4 and 15.0 g agar.
Biochemical tests were carried out using API 20NE, API
20E, API ID32 GN and API ZYM test kits (bioMérieux) as
described by the manufacturer except that the cultures were
suspended in ASW. Growth was assessed at temperatures of
4, 7, 15, 28 and 30 uC, and from 32 to 40 uC in increments of
1 uC. Antibiotic susceptibility was examined using commercial paper discs (Research Centre of Pharmacotherapy, St.
Petersburg) impregnated with the following antibiotics (mg
per disc, unless otherwise indicated): ampicillin (10),
benzylpenicillin (10 U), vancomycin (30), gentamicin (10),
kanamycin (30), carbenicillin (100), chloramphenicol (30),
neomycin (30), oxacillin (10), oleandomycin (15), lincomycin (15), ofloxacin (5), rifampicin (5), polymyxin
(300 U), streptomycin (30), cephazolin (30), cephalexin
(30), erythromycin (15), nalidixic acid (30), tetracycline (30)
and doxycycline (10 g). The pH range for growth was
determined in MB adjusted to pH 5.0–11.5 (in increments
of 0.5 pH unit) by the addition of 1 M HCl or 1 M NaOH.
For polar lipid and fatty acid analyses, strains KMM 9530T,
L. sediminilitoris KCTC 32383T, L. maricola JCM 14564T, L.
tamlensis JCM 14020T and L. rosea KMM 6003T were grown
on MA at 28 uC for 3 days. Lipids were extracted using the
method of Folch et al. (1957). Two-dimensional TLC of
polar lipids was carried out on Silica gel 60 F254 plates
(10610 cm; Merck) using chloroform/methanol/water
(65 : 25 : 4, by vol.) for the first direction, and chloroform/
methanol/acetic acid/water (80 : 12 : 15 : 4, by vol.) for the
second direction (Collins & Shah, 1984). Non-specific
detection of lipids by two-dimensional TLC was achieved
with 10 % H2SO4 in methanol at 120 uC. Amino-containing
lipids were determined with ninhydrin, phospholipids with
http://ijs.sgmjournals.org
molybdate reagent, glycolipids with a-naphthol, and cholinecontaining lipids with Dragendorff’s reagent. Respiratory
lipoquinones were analysed by reversed-phase HPLC as
described by Mitchell & Fallon (1990). Fatty acid methyl
esters (FAMEs) were prepared according to the procedure of
the Microbial Identification System (MIDI) (Sasser, 1990).
The analysis of FAMEs was performed as described
previously (Romanenko et al., 2011a). Production of
bacteriochlorophyll a was tested spectrophotometrically in
methanolic extracts of cells grown on MA and MB in the
dark as described by Lafay et al. (1995). The 16S rRNA gene
sequence (1447 nt) of strain KMM 9530T was determined as
described by Shida et al. (1997). The sequence obtained was
compared with 16S rRNA gene sequences retrieved from the
EMBL/GenBank/DDBJ databases by using the FASTA program (Pearson & Lipman, 1988). Phylogenetic analysis of
16S rRNA gene sequences was performed using the software
package MEGA 5 (Tamura et al., 2011) after multiple
alignment of data by CLUSTAL W (version 1.83; Thompson
et al., 2002). Phylogenetic trees were reconstructed by the
neighbour-joining, maximum-likelihood and maximumparsimony methods and distances were calculated according
to the Kimura two-parameter model. The robustness of
phylogenetic trees was estimated by the bootstrap analysis of
1000 replicates. DNA–DNA hybridization experiments were
performed by the photobiotin-labelled DNA probe microplate method of Ezaki et al. (1989). DNA–DNA hybridization values are given as means of at least two determinations.
Comparative 16S rRNA gene sequence analysis established
that strain KMM 9530T belonged to the genus Loktanella,
displaying the highest sequence similarity values of 98.5,
98.4 and 98.2 % to the closely related species of the genus
Loktanella, L. sediminilitoris KCTC 32383T, L. tamlensis
JCM 14020T and L. maricola JCM 14564T, respectively, and
similarity values below 97.7 % to the remaining species of
the genus Loktanella (Fig. 1). The 16S rRNA gene sequence
similarities obtained for strain KMM 9530T and most species
of the genus Loktanella were lower than the threshold
similarity value of 97 % proposed by Stackebrandt & Goebel
(1994) and re-evaluated to 98.7 % by Stackebrandt & Ebers
(2006), indicating that the novel strain could be assigned to
the genus Loktanella as an individual species. DNA–DNA
hybridization experiments were carried out between strain
KMM 9530T and the type strains L. sediminilitoris KCTC
32383T, L. tamlensis JCM 14020T and L. maricola JCM
14564T, and DNA relatedness values were determined to be
22, 45 and 36 %, respectively. The DNA–DNA hybridization
values confirmed the assignment of the novel isolate KMM
9530T to a separate species according to the cut-off value of
70 % proposed by Wayne et al. (1987) for the bacterial
species delineation.
Cells of strain KMM 9530T were aerobic, Gram-stainnegative, non-motile ovoid or short rod-shaped (Fig. S1,
available in the online Supplementary Material). Cultural,
physiological and metabolic properties of strain KMM
9530T and related members of the genus Loktanella are
listed in Table 1 and in the species description. Fatty acid
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N. Tanaka and others
*/*/65
76/*/83
92/*/*
Loktanella tamlensis JCM 14020T (DQ533556)
Loktanella maritima KMM 9530T (AB894236)
Loktanella sediminilitoris KCTC 32383T (KC311338)
Loktanella maricola JCM 14564T (EF202613)
82/*/*
Loktanella rosea KMM 6003T (AY682199)
Loktanella koreensis GA2-M3T (DQ344498)
Loktanella litorea DPG-5T (JN885197)
Loktanella agnita R10SW5T (AY682198)
Loktanella vestfoldensis LMG 22003T (AJ582226)
0.01
63/*/*
100/*/100
Loktanella salsilacus LMG 21507T (AJ440997)
Loktanella atrilutea IG8T (AB246747)
Loktanella fryxellensis LMG 22007T (AJ582225)
Loktanella pyoseonensis JJM85T (AM983542)
79/65/74
100/100/99
Loktanella hongkongensis UST950701-009PT (AY600300)
Loktanella cinnabarina LL-001T (AB688112)
Sagittula stellata E-37T (U58356)
Staleya guttiformis EL-38T (Y16427)
86/70/83
90/67/89
Sulfitobacter pontiacus ChLG 10T (Y13155)
Jannaschia helgolandensis Hel 10T (AJ438157)
96/90/95
Thalassobacter stenotrophicus CECT 5294T (AJ631302)
Leisingera methylohalidivorans MB2T (AY005463)
Ketogulonicigenium vulgare DSM 4025T (AF136849)
Stappia stellulata IAM 12621T (D88525)
Fig. 1. Neighbour-joining tree based on 16S rRNA gene sequences showing relationships of isolate KMM 9530T and related
taxa. Filled circles, generic branches that are present in phylogenetic trees generated by the neighbour-joining, maximumlikelihood and maximum-parsimony methods; filled triangles, generic branches that are present in trees generated by the
neighbour-joining and maximum-parsimony methods; filled squares, generic branches that are present in trees generated by the
neighbour-joining and maximum-likelihood methods. Numbers indicate bootstrap values as percentages greater than 60
(neighbour-joining probability/maximum-parsimony probability/maximum-likelihood probability). These values are based on
1000 replicates. Bar, 0.01 substitutions per nucleotide position.
profiles were similar with a large proportion of C18 : 1v7c
and the presence of 11-methyl C18 : 1v7c found in all strains
tested (Table 2). These results are in accordance with the
data reported for L. rosea KMM 6003T (Ivanova et al.,
2005) and L. maricola JCM 14564T (Yoon et al., 2007), but
disagreed with the results of Lee (2012) and Park et al.
(2013a) who did not find 11-methyl C18 : 1v7c in L.
tamlensis JCM 14020T and L. sediminilitoris KCTC 32383T.
Strains KMM 9530T and L. tamlensis JCM 14020T differed
from the other related strains by a smaller proportion of
11-methyl C18 : 1v7c, and only L. sediminilitoris KCTC
32383T contained C12 : 1 (Table 2). The major isoprenoid
quinone of strain KMM 9530T was ubiquinone Q-10.
The polar lipid composition of strain KMM 9530T and
related strains L. tamlensis JCM 14020T, L. rosea KMM
6003T, L. maricola JCM 14564T, L. sediminilitoris KCTC
32383T was found to be similar and included phosphatidylcholine (PC), phosphatidylglycerol (PG), diphosphatidylglycerol (DPG), an unknown aminolipid, unknown
phospholipids, and unknown lipids. All strains tested
contained minor amounts of DPG (Fig. S2). The polar
profile of L. sediminilitoris KCTC 32383T differed from
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those of the other four strains studied in that phosphatidylethanolamine (PE) was detected. This finding is
congruent with the original description by Park et al.
(2013a) who reported the presence of PC, PG, PE and an
unknown aminolipid as dominant and DPG, an unknown
phospholipid and an unknown lipid as minor components
in L. sediminilitoris KCTC 32383T. In addition, unknown
phospholipids PL1–PL3 were found in the present study.
The presence of PE in L. sediminilitoris KCTC 32383T and
its absence in KMM 9530T and the three related strains of
the genus Loktanella was confirmed by two-dimensional
TLC, spraying with ninhydrin followed by staining with
molybdate reagent, as exemplified by strains KMM 9530T,
L. maricola JCM 14564T and L. sediminilitoris KCTC
32383T in Fig. S2. Unlike our results with regard to the lack
of PE in the three above type strains, the occurrence of PE
has been reported by Lee (2012), Yoon et al. (2007) and
Ivanova et al. (2005) in L. tamlensis JCM 14020T, L.
maricola JCM 14564T and L. rosea KMM 6003T, with
minor and trace amounts for the two latter strains. The
genus Loktanella comprises both bacteria having PE
together with PC and PG, L. litorea (Yoon et al., 2013),
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Loktanella maritima sp. nov. from shallow marine sediments
Table 1. Differential phenotypic characteristics of strain KMM 9530T and type strains of related species of the genus Loktanella
Strains: 1, Loktanella maritima sp. nov. KMM 9530T; 2, L. tamlensis JCM 14020T; 3, L. maricola JCM 14564T; 4, L. rosea KMM 6003T; 5, L.
sediminilitoris KCTC 32383T. All data were obtained from the present study. +, Positive; 2, negative; (+), weakly positive reaction. All strains are
positive for oxidase, catalase, sodium ion requirement for growth, aesculin hydrolysis and p-nitrophenyl-b-D-galactopyranoside (PNPG) test
(b-galactosidase) (in API 20NE), and production of esterase (C4) and esterase lipase (C8). All strains are negative for gelatin hydrolysis and nitrate
reduction (in routine and API 20NE tests), hydrolysis of casein, starch*, chitin and xanthine, production of lipase (C14), cystine arylamidase,
a-chymotrypsin, a-galactosidase, b-glucuronidase, N-acetyl-b-glucosaminidase, b-galactosidase, a-mannosidase, a-fucosidase (in API ZYM),
arginine dihydrolase and urease, glucose fermentation, indole production, and assimilation of D-glucose, L-arabinose, D-mannose, D-mannitol,
N-acetylglucosamine, maltose, D-gluconate, caprate, adipate, L-malate, citrate and phenylacetate (in API 20NE). All strains are susceptible to
ampicillin, benzylpenicillin, vancomycin, gentamicin, kanamycin, carbenicillin, chloramphenicol, streptomycin, oleandomycin, ofloxacin,
neomycin, oxacillin, rifampicin, cephazolin, cephalexin and erythromycin, and resistant to polymyxinD.
Characteristic
Colony pigmentation
Motility
Temperature range for growth (6C)
NaCl range for growth (%)
Hydrolysis of:
Tyrosine
Hypoxanthine
Tween 80
DNA
Citrate utilization (Simmons’ citrate agar)
H2S formation
API ZYM tests
Alkaline phosphatase
Leucine arylamidase
Valine arylamidase
Trypsin
Acid phosphatase
Naphthol-AS-BI-phosphohydrolase
a-Glucosidase
b-Glucosidase
Sensitivity to antibiotics (mg/disc)
Lincomycin (15)
Nalidixic acid (30)
Tetracycline (30)
Doxycycline (10)
1
2
3
4
5
Beige
”
4–36
2–8
Beige
+
4–34*
2–4*
Dark pinkd
”
4–34
3–6d
Pink
+
4–39§
2–8§
Greyish-yellow
”
4–35D
2–5D
”
(+)
”
”
+
”
”
+
”
”
”
”
”
+
”
”
”
”
+
+
”
”
”
”
”
”
+
+
”
+
+
+
(+)
”
+
+
(+)
+
+
+
”
”
+
+
”
”
+
+
”
”
+
+
”
”
(+)
+
”
”
(+)
+
”
”
+
”
”
+
”
(+)D
”
”
”
”
”
”
+
+
+
+
+
+
+
+
”
+
+
+
”
”
+
+
*Results differ from those reported by Lee (2012).
DResults differ from those reported by Park et al. (2013a).
dData not consistent with those reported by Yoon et al. (2007). Colony pigmentation for L. maricola JCM 14564T was described by Yoon et al.
(2007) as light orange.
§Results differ from those reported by Ivanova et al. (2005).
L. agnita (Ivanova et al., 2005), L. sediminilitoris KCTC
32383T (Park et al., 2013a) and L. cinnabarina (Tsubouchi
et al., 2013), and bacteria not containing PE in their polar
lipid profiles, L. pyoseonensis (Moon et al., 2010), L. hongkongensis (Tsubouchi et al., 2013) and L. soesokkakensis
(Park et al., 2013b). At the same time, Yoon et al. (2013)
reported the presence of PE in L. salsilacus whereas PE was
not found in this bacterium in the study of Tsubouchi et al.
(2013). Tsubouchi et al. (2013) and Park et al. (2013b)
reported that the same L. cinnabarina cluster includes
bacteria containing PE (L. cinnabarina itself) and bacteria
without PE (L. pyoseonensis, L. hongkongensis and L.
http://ijs.sgmjournals.org
soesokkakensis). Chemotaxonomic properties (ubiquinone
Q-10, the predominance of C18 : 1v7c, and the presence of
PC and PG) obtained for strain KMM 9530T supported its
assignment to the genus Loktanella.
It is evident from the results obtained that the novel isolate
can be assigned to the genus Loktanella on the basis of its
physiological, biochemical and chemotaxonomic characteristics. It is interesting to note that type strains of species
of the genus Loktanella revealed similar antibiotic resistance profiles, being susceptible to 19 or 20 of the 21
antibiotics applied, except for strains KMM 9530T and L.
sediminilitoris KCTC 32383T. Recently we have reported
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N. Tanaka and others
Table 2. Fatty acid (%) profiles of strain KMM 9530T and
related members of the genus Loktanella
Strains: 1, Loktanella maritima sp. nov. KMM 9530T; 2, L. tamlensis
JCM 14020T; 3, L. maricola JCM 14564T; 4, L. rosea KMM 6003T; 5, L.
sediminilitoris KCTC 32383T. Results were obtained from present
study. All strains were grown on MA at 28 uC for 3 days. –, Not
detected; TR, trace amount.
Fatty acid
C10 : 03-OH
C12 : 13-OH
C12 : 1
C16 : 1v7c
C16 : 0
C17 : 1v8c
C17 : 0
C18 : 2
C18 : 1v7c
11-Methyl C18 : 1v7c
C18 : 0
1
2
3
4
5
3.09
2.45
–
0.49
5.76
1.07
2.28
3.40
76.98
2.62
1.40
3.29
1.95
–
1.92
6.98
–
0.55
1.65
75.21
4.56
1.25
3.43
4.37
–
0.39
5.34
–
0.50
1.45
68.62
9.70
2.20
1.09
3.53
1.48
3.37
5.29
1.12
12.45
–
0.39
4.69
61.96
6.78
3.95
TR
1.09
6.32
0.26
0.89
3.69
68.02
11.47
1.96
marine alphaproteobacteria showing a high sensitivity to
antibiotics (Romanenko et al., 2011a, b). Strain KMM
9530T could be distinguished from related type strains of
species of the genus Loktanella by colony pigmentation
(except for L. tamlensis JCM 14020T), being able to grow
with 7–8 % NaCl (except for L. rosea KMM 6003T) and
being able to utilize citrate, as well as by enzyme activities
in API ZYM tests and its antibiotic sensitivity profile. It
should be noted that the recently described L. sediminilitoris
KCTC 32383T revealed a number of distinctive traits
compared with related type strains of species of the genus
Loktanella tested, including hydrolysis of Tween 80 and
DNA and API ZYM enzyme activity profile. Differential
phenotypic characteristics are listed in Table 1. On the basis
of the results obtained, it is proposed to assign strain KMM
9530T to the genus Loktanella as a representative of a novel
species, Loktanella maritima sp. nov.
Description of Loktanella maritima sp. nov.
Loktanella maritima (ma.ri9ti.ma. L. fem. adj. maritima
maritime, marine).
Gram-stain-negative, aerobic, oxidase-positive, catalasepositive, ovoid or short rod-shaped, non-motile cells, 0.6–
0.8 mm in diameter and 1.6–2.0 mm in length. Grows on
MA 2216 and MB. Produces light beige-pigmented, shiny
smooth colonies with regular edges, 2–3 mm in diameter.
No growth is observed on commercial tryptic soy agar or
broth, nutrient agar or R2A agar. Bacteriochlorophyll a is
not produced. Requires NaCl for growth; growth occurs
with between 2 and 8 % (w/v) NaCl with an optimum of 3–
4 % NaCl; growth is weak with 2 and 8 % NaCl. The
temperature range for growth is 4–36 uC with an optimum
of 28–30 uC; growth is weak at 36 uC. The pH range for
growth is pH 5.5–9.0 (optimal growth at pH 6.5–7.5).
2374
Negative for hydrolysis of casein, DNA, chitin, starch,
Tween 80, L-tyrosine and xanthine, and production of H2S
in conventional tests. Negative for gelatin hydrolysis and
nitrate reduction in routine and API 20NE tests. Positive
for aesculin hydrolysis (API 20NE), p-nitrophenyl-b-Dgalactopyranoside (PNPG) test (b-galactosidase in API
20NE and API 20E tests) and citrate utilization (API 20E);
negative for the remaining tests that are included to the
API 20NE and API 20E panels. Cannot assimilate any of
the substrates included in the ID32 GN gallery. Positive API
ZYM test results for alkaline phosphatase, esterase (C4),
esterase lipase (C8), leucine arylamidase, acid phosphatase,
naphthol-AS-BI-phosphohydrolase and b-glucosidase; weakly
positive for valine arylamidase and a-glucosidase; and
negative for lipase (C14), cystine arylamidase, trypsin, achymotrypsin, a-galactosidase, b-galactosidase, b-glucuronidase, N-acetyl-b-glucosaminidase, a-mannosidase and
a-fucosidase. The major isoprenoid quinone is ubiquinone
Q-10. The polar lipids include PC, PG, DPG, an unknown
aminolipid, an unknown phospholipid and four unknown
lipids. Fatty acid C18 : 1v7c is predominant. Susceptible to
ampicillin, benzylpenicillin, vancomycin, gentamicin, kanamycin, carbenicillin, chloramphenicol, neomycin, oxacillin,
oleandomycin, ofloxacin, rifampicin, streptomycin, cephazolin, cephalexin and erythromycin; and resistant to
lincomycin, polymyxin, nalidixic acid, tetracycline and
doxycycline.
The type strain is KMM 9530T (5NRIC 0919T5JCM
19807T), isolated from a shallow sediment sample,
collected from Peter the Great Bay, the Sea of Japan,
Russia.
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