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Int J Syst Evol Microbiol 54 (2004), 1017-1023; DOI  10.1099/ijs.0.02849-0
© 2004 International Union of Microbiological Societies

Maribacter gen. nov., a new member of the family Flavobacteriaceae, isolated from marine habitats, containing the species Maribacter sedimenticola sp. nov., Maribacter aquivivus sp. nov., Maribacter orientalis sp. nov. and Maribacter ulvicola sp. nov.

Olga I. Nedashkovskaya1, Seung Bum Kim2, Suk Kyun Han2, Anatoly M. Lysenko3, Manfred Rohde4, Moon-Soo Rhee2, Galina M. Frolova1, Enevold Falsen5, Valery V. Mikhailov1 and Kyung Sook Bae2

1 Pacific Institute of Bioorganic Chemistry of the Far-Eastern Branch of the Russian Academy of Sciences, Pr. 100 Let Vladivostoku 159, 690022, Vladivostok, Russia
2 Korean Collection for Type Cultures, Korea Institute of Bioscience and Biotechnology, Yusong, Daejon 305-333, Republic of Korea
3 Institute of Microbiology of the Russian Academy of Sciences, Pr. 60 let October 7/2, Moscow, 117811, Russia
4 Bereich Mikrobiologie, Abt. Mikrobielle Pathogenitat und Impfstoffforschung, GBF – Gesellschaft für Biotechnologische Forschung, Mascheroder Weg 1, D-38124 Braunschweig, Germany
5 Culture Collection, Department of Clinical Bacteriology, University of Göteborg, Guldhedsgatan 10, S-413 46 Göteborg, Sweden

Correspondence
Olga I. Nedashkovskaya
olganedashkovska{at}piboc.dvo.ru
or
olganedashkovska{at}yahoo.com


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Six novel gliding, heterotrophic, Gram-negative, yellow-pigmented, aerobic, oxidase- and catalase-positive bacteria were isolated from the green alga Ulva fenestrata, sea water and a bottom sediment sample collected in the Gulf of Peter the Great, Sea of Japan. 16S rRNA gene sequence analysis revealed that the strains studied were members of the family Flavobacteriaceae. On the basis of their phenotypic, chemotaxonomic, genotypic and phylogenetic characteristics, the novel bacteria have been assigned to the new genus Maribacter gen. nov., as Maribacter sedimenticola sp. nov., Maribacter orientalis sp. nov., Maribacter aquivivus sp. nov. and Maribacter ulvicola sp. nov., with the type strains KMM 3903T (=KCTC 12966T=CCUG 47098T), KMM 3947T (=KCTC 12967T=CCUG 48008T), KMM 3949T (=KCTC 12968T=CCUG 48009T) and KMM 3951T (=KCTC 12969T=DSM 15366T), respectively.

Published online ahead of print on 17 October 2003 as DOI 10.1099/ijs.0.02849-0.

The GenBank/EMBL/DDBJ accession numbers for Maribacter sedimenticola KMM 3903T, Maribacter orientalis KMM 3947T, Maribacter aquivivus KMM 3949T and Maribacter ulvicola KMM 3951T are AY271623, AY271624, AY271625 and AY271626, respectively.

Micrographs of some of the strains described in this study are available from IJSEM Online.


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
The gliding agarolytic bacteria belonging to the family Flavobacteriaceae are often isolated from different marine environments. Many novel bacteria that decompose agar, a polysaccharide of the red alga, have been collected from coastal bottom sediments. For example, a single strain of Zobellia uliginosa ATCC 14397T, formerly Flavobacterium uliginosum (ZoBell & Upham, 1944Go), was found in a beach mud sample collected near California (Barbeyron et al., 2001Go). A common coastal dweller, Cellulophaga lytica, formerly [Cytophaga] lytica (Reichenbach, 1989Go), was originally isolated from beach sand in Costa Rica (Lewin, 1969Go; Johansen et al., 1999Go). Some of the agar-degrading bacteria are associated with algae. For example, two species of the genus Cellulophaga, Cellulophaga baltica and Cellulophaga fucicola, were isolated from brown alga Fucus serratus collected in the North Sea (Johansen et al., 1999Go). The diatom Melosira sp. and an unidentified macrophyte have been colonized by representatives of another species of the genus Cellulophaga, Cellulophaga algicola (Bowman, 2000Go). The type strain of the type species of the genus Zobellia, Zobellia galactanivorans DsijT, was isolated from surfaces of the red alga Delesseria sanguinea (Barbeyron et al., 2001Go).

In this study, we describe the phenotypic and genotypic characteristics of six gliding, Gram-negative bacteria isolated from the green alga Ulva fenestrata, sea water and a bottom sediment collected in the Gulf of Peter the Great, Sea of Japan. A 16S rRNA gene-based phylogenetic analysis revealed that the strains studied should be assigned to a new genus, Maribacter gen. nov., as four species within the Muricauda–Zobellia–Arenibacter cluster of the family Flavobacteriaceae. The names Maribacter aquivivus sp. nov., Maribacter orientalis sp. nov., Maribacter sedimenticola sp. nov. and Maribacter ulvicola sp. nov. are proposed for the novel marine bacteria.


   METHODS
TOP
 ABSTRACT
 INTRODUCTION
 METHODS
RESULTS AND DISCUSSION
 REFERENCES
 
Organisms.
Strains KMM 3947T, KMM 3948 and KMM 3949T were isolated from sea water; KMM 3903T was isolated from a bottom sediment sample; KMM 3951T and KMM 3952 were isolated from the green alga Ulva fenestrata collected in the Gulf of Peter the Great, Sea of Japan, during June 2000. Strains were cultivated at 28 °C on marine agar (Difco) and stored at –80 °C in marine broth (Difco) supplemented with 20 % (v/v) glycerol.

Phenotypic characterization.
Oxidative or fermentative utilization of glucose was determined using the Hugh–Leifson medium modified for marine bacteria (Lemos et al., 1985Go). Degradation of agar, starch, casein, gelatin, cellulose (filter paper and CMC), chitin, DNA, urea and alginic acids, flexirubin pigment production, growth at different NaCl concentrations or pH, production of acid from carbohydrates and susceptibility to antibiotics were tested as described previously (Nedashkovskaya et al., 2003aGo, bGo). Gram-staining reaction; hydrolysis of Tween 20, Tween 40 and Tween 80; nitrate reduction; production of hydrogen sulphide and indole; and {beta}-galactosidase, oxidase, catalase and alkaline phosphatase activities were tested according to the methods of Gerhardt et al. (1994)Go. To examine carbon source utilization, a medium containing 0·2 g NaNO3, 0·2 g NH4Cl, 0·05 g yeast extract (Difco) and 0·4 % (w/v) carbon source in 1 litre of artificial sea water and the commercial API 20NE identification strips, following the instructions of the manufacturers, were used. To study the growth temperature ranges of the strains, bacteria were cultivated on medium A consisting of (l–1) 5 g Bacto peptone (Difco), 2 g Bacto yeast extract (Difco), 1 g glucose, 0·02 g KH2PO4 and 0·05 g MgSO4.7H2O in 50 % (v/v) natural sea water and 50 % (v/v) distilled water. Spread growth was observed by cultivation on medium B which contained (l–1) 1 g Bacto peptone (Difco), 1 g yeast extract (Difco), 15 g agar and half-strength natural sea water under high-moisture conditions. Gliding motility was determined as described by Bowman (2000)Go. The cell movement at the edges of colonies was verified by using phase-contrast microscopy.

Scanning electron microscopy.
The bacteria were fixed with a solution containing 2 % glutaraldehyde and 3 % formaldehyde in cacodylate buffer (0·1 M cacodylate, 0·09 M sucrose, 0·01 M CaCl2, 0·01 M MgCl2, pH 6·9) for 1 h on ice and washed with cacodylate buffer. After being washed several times in TE buffer (20 mM Tris, 1 mM EDTA, pH 7·0), samples were dehydrated with a graded series of acetone (10, 30, 50, 70, 90, 100 %) on ice, each step 15 min, followed by critical-point drying with liquid CO2. Samples were sputter-coated with an ~10 nm thick gold film before being examined in a Zeiss field emission scanning electron microscope DSM982 Gemini at an acceleration voltage of 5 kV using the Everhart Thornley SE detector and the inlens-SE detector in a 50 : 50 ratio.

Cellular fatty acid composition and respiratory quinone analyses.
The analysis of fatty acid methyl esters was carried out according to the standard protocol of the Microbial Identification System (Microbial ID). Isoprenoid quinones were extracted from lyophilized cells and analysed as described by Akagawa-Matsushita et al. (1992)Go. Isoprenoid quinone composition was characterized by HPLC (Shimadzu instruments) using a reversed-phase type Zorbax ODS column (250x4·6 mm) and acetonitrile/2-propanol (65 : 35, v/v) as a mobile phase at a flow rate of 0·5 ml min–1. The column was kept at 40 °C. Menaquinones were detected by monitoring at 270 nm and were identified by comparison with known quinones from a reference strain, Salegentibacter salegens DSM 5424T.

DNA base content and DNA–DNA reassociation.
DNA was isolated following the method of Marmur (1961)Go and the G+C content (mol%) of the DNA was determined by the thermal denaturation method (Marmur & Doty, 1962Go). DNA–DNA hybridization was performed spectrophotometrically and initial renaturation rates were recorded as described by De Ley et al. (1970)Go.

16S rRNA gene sequencing and phylogenetic analysis.
DNA extraction, PCR and sequencing of 16S rRNA gene followed previously described procedures (Kim et al., 1998Go). The obtained sequence data were aligned together with those of representative members of selected genera belonging to the family Flavobacteriaceae using PHYDIT version 3.2 (http://plaza.snu.ac.kr/~jchun/phydit/). Phylogenetic trees were inferred using suitable programs of the PHYLIP package (Felsenstein, 1993Go). Phylogenetic distances were calculated from the models of Jukes & Cantor (1969)Go, and the trees were constructed on the basis of the neighbour-joining (Saitou & Nei, 1987Go), least squares (Fitch & Margoliash, 1967Go) and maximum-likelihood (Felsenstein, 1993Go) algorithms. Bootstrap analysis was performed with 1000 resampled data sets, using SEQBOOT and CONSENSE programs of the PHYLIP package.


   RESULTS AND DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS AND DISCUSSION
REFERENCES
 
Bacteria isolated in this study were gliding, Gram-negative, chemo-organotrophic organisms with a respiratory type of metabolism. They were non-motile, single, flexible rods, of 0·5–0·7 µm in diameter and 2–10 µm in length (micrographs, supplementary data in IJSEM Online). On marine agar, colonies of strains studied were round, yellow-pigmented, shiny, with entire edges, and were 2–4 mm in diameter. Colonies of strains KMM 3903T, KMM 3948, KMM 3949T and KMM 3951T were weakly sunken into the agar. The bacteria studied were oxidase-, catalase- and alkaline phosphatase-positive, and required Na+ ions for growth. Growth of strains KMM 3947T, KMM 3951T and KMM 3952 occurred in the presence of 1–5 % NaCl, strain KMM 3903T grew in media containing 1–6 % NaCl and strain KMM 3949T grew in the presence of 1–7 % NaCl; optimal growth for all strains was observed at 1·5–2·0 % NaCl. All strains grew at 4 °C. The maximum temperatures for growth of strains KMM 3903T, KMM 3947T, KMM 3949T and KMM 3951T were 33, 32, 30 and 32 °C, respectively. The pH range for growth was 5·5–10·0, with optimum growth occurring between pH 7·5 and 8·5. Other physiological and biochemical characteristics of the KMM strains described in this study are listed in Table 1Go. Predominant cellular fatty acids of strains studied were of the branched-chain unsaturated and straight-chain saturated types, namely i-C15 : 0, i-C15 : 1, C15 : 0 and i-C17 : 03-OH fatty acids (Table 2Go). The main respiratory quinone was MK-6. The DNA base contents of strains KMM 3903T, KMM 3947T, KMM 3949T, KMM 3951T and KMM 3952 were 37·5, 39·0, 35·0, 36·7 and 35·9 mol%, respectively, as determined by the thermal denaturation method. The level of DNA–DNA reassociation between strains KMM 3948 and KMM 3949T, KMM 3951T and KMM 3952, and KMM 3949T and KMM 3951T was 85, 93 and 51 %, respectively.


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  		Table 1. Phenotypic characteristics of Maribacter species

Species: 1, M. sedimenticola KMM 3903T; 2, M. orientalis KMM 3947T; 3, M. aquivivus KMM 3949T; 4, M. ulvicola KMM 3951T. All strains were positive for: respiratory metabolism; gliding motility; oxidase, catalase and alkaline phosphatase activity; NaCl requirement for growth; growth at 4–30 °C and in the presence of 1–5 % NaCl; hydrolysis of Tween 40; susceptibility to carbenicillin, lincomycin and oleandomycin. All strains were negative for: hydrolysis of casein, cellulose (CMC, filter paper), chitin and urea; production of flexirubin pigments; acid formation from raffinose, sorbose, N-acetylglucosamine, citrate, adonitol, glycerol, inositol, mannitol and sorbitol; utilization of mannitol, inositol, sorbitol, malonate and citrate; indole, H2S, acetoin (Voges–Proskauer reaction); susceptibility to benzylpenicillin, gentamicin, kanamycin, neomycin, polymyxin B and streptomycin. +, Positive test result; –, negative test result.

 

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  		Table 2. Whole-cell fatty acid profiles (percentage composition) of Maribacter species

Species: 1, M. sedimenticola KMM 3903T; 2, M. aquivivus KMM 3949T; 3, M. ulvicola KMM 3951T; 4, M. orientalis KMM 3947T.

 
A phylogenetic analysis of almost-complete 16S rRNA gene sequences revealed that the strains studied formed a distinct lineage within the phylum Cytophaga–Flavobacterium–Bacteroides and created a cluster with Arenibacter latericius, Muricauda ruestringensis and Zobellia species (Fig. 1Go). The sequence identities of KMM strains with Arenibacter latericius and Muricauda ruestringensis were 90·9–91·5 % and 89·9–90·4 %, respectively. The 16S rRNA gene sequence similarity values for KMM strains and their nearest neighbour, Z. uliginosa, were 92·9, 93·1, 94·3 and 93·8 %, respectively, but their relationships were without significant bootstrap support. Differential features of strains KMM 3903T, KMM 3947T, KMM 3949T and KMM 3951T and related members of the phylum Cytophaga–Flavobacterium–Bacteroides are presented in Table 3Go. The production of flexirubin-type pigments, casein hydrolysis and maximum growth temperature (42–43 °C) differentiate members of the genus Zobellia from the strains studied. The low level of sequence similarity of the strains tested with other Cytophaga-like bacteria described to date (84·5–91·6 %) clearly demonstrates that the bacteria isolated in this study represent a new genus. The 16S rRNA gene sequences of the novel strains showed between 95·1 and 97·4 % similarity; only the 16S rRNA gene sequence similarity value between strains KMM 3949T and KMM 3951T was higher than 97 %. It is known that strains which demonstrate less than 3 % difference in their 16S rRNA gene sequences may belong to the same species (Stackebrandt & Goebel, 1994Go). The DNA–DNA hybridization experiment carried out between strains KMM 3949T and KMM 3951T showed a DNA–DNA reassociation value of 51 %. Consequently, strains KMM 3949T and KMM 3951T were shown to be separate species.



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  		Fig. 1. Phylogenetic tree based on the 16S rRNA gene sequences of the KMM strains described in this study and representative members of related genera in the family Flavobacteriaceae. The asterisks indicate branches that were also recovered using least-squares and maximum-likelihood algorithms. The numbers at nodes indicate bootstrap values (%). Bar, 0·01 substitutions per nucleotide position.

 

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  		Table 3. Characteristics that differentiate the genus Maribacter from other, related, genera of the family Flavobacteriaceae

Genus: 1, Maribacter; 2, Zobellia; 3, Arenibacter; 4, Muricauda; 5, Vitellibacter; 6, Aequorivita; 7, Cellulophaga; 8, [Cytophaga] marinoflava. Data are from Reichenbach (1989)Go, Johansen et al. (1999)Go, Bowman (2000)Go, Ivanova et al. (2001)Go, Barbeyron et al. (2001)Go, Bruns et al. (2001)Go, Bowman & Nichols (2002)Go, Nedashkovskaya et al. (2003aGo, b)Go and this study. A, Aerobic; F, facultative; +, positive test result; –, negative test result; V, variable; ND, not determined.

 
Thus, sequence distinctiveness and DNA–DNA reassociation data define strains KMM 3903T, KMM 3947T, KMM 3949T and KMM 3951T as separate genospecies. Phenotypic findings support the data from the genomic analyses of the strains studied (Table 1Go). Physiologically, strain KMM 3903T may be differentiated from other strains studied by the absence of {beta}-galactosidase activity, non-utilization of the majority of carbohydrates tested, growth at 37 °C, starch hydrolysis and susceptibility to tetracycline; strain KMM 3947T does not hydrolyse Tween 80, but forms acid from galactose, xylose and melibiose; strain KMM 3949T produces DNase, does not hydrolyse Tween 20 and grows at 30 °C. Also, strain KMM 3951T may be easily separated from the above strains by the absence of alginate hydrolysis, fucose and rhamnose oxidation and susceptibility to ampicillin.

Thus, our conclusion, supported by the polyphasic analysis data presented in this report, is that the bacteria studied here could not be assigned to any of the taxa currently included in the phylum Cytophaga–Flavobacterium–Bacteroides. Consequently, we propose that strains KMM 3903T, KMM 3947T, KMM 3949T and KMM 3948, and KMM 3951T and KMM 3952 be placed in a new genus, Maribacter, as Maribacter sedimenticola, Maribacter orientalis, Maribacter aquivivus and Maribacter ulvicola, respectively.

Description of Maribacter gen. nov.
Maribacter (Ma.ri.bac'ter. L. neut. n. mare the sea; N.L. masc. n. bacter from Gr. n. bakteron rod; N.L. masc. n. Maribacter rod inhabiting marine environments).

Rod-shaped cells. Non-motile. Gram-negative. Do not form endospores. Require Na+ ions for growth. Strictly aerobic. Produce non-diffusible yellow pigments. No flexirubins are formed. Chemo-organotroph. Cytochrome oxidase-, catalase- and alkaline phosphatase-positive. Predominant cellular fatty acids are straight-chain saturated, branched-chain saturated and unsaturated fatty acids i-C15 : 0, i-C15 : 1, C15 : 0 and i-C17 : 03-OH. The main lipoquinone is MK-6. As determined by 16S rRNA gene sequence analysis, the genus Maribacter is a member of the family Flavobacteriaceae, phylum Cytophaga–Flavobacterium–Bacteroides.

Type species is Maribacter sedimenticola.

Description of Maribacter sedimenticola sp. nov.
Maribacter sedimenticola (se.di.men.ti.co'la. L. masc. n. sedimentum sediment; L. suffix -cola dweller; N.L. masc. n. sedimenticola sediment dweller).

Main characteristics are the same as those given for the genus. In addition, cells range in size from 0·5 to 0·7 µm in width and 2 to 10 µm in length. On marine agar, colonies are 2–4 mm in diameter, circular, shiny with entire edges, yellow pigmented and weakly sunken into agar. Grows at 4–33 °C; optimal temperature for growth is between 22 and 24 °C. Grows in the presence of 1–6 % NaCl. Decomposes agar, gelatin, alginate, DNA, Tween 40 and Tween 80. Does not hydrolyse casein, starch, Tween 20, cellulose (carboximethylcellulose and filter paper) or chitin. Forms no acid from cellobiose, fucose, galactose, glucose, lactose, maltose, melibiose, raffinose, rhamnose, sucrose, xylose, citrate, adonitol, dulcitol, glycerol, inositol or mannitol. Does not utilize glucose, lactose, mannose, sucrose, mannitol, inositol, sorbitol, malonate or citrate. Nitrate is not reduced. H2S, indole and acetoin (Voges–Proskauer reaction) production are negative. Susceptible to carbenicillin, lincomycin, oleandomycin and tetracycline; resistant to ampicillin, benzylpenicillin, gentamicin, kanamycin, neomycin, polymyxin B and streptomycin.

Type strain is KMM 3903T (=KCTC 12966T=CCUG 47098T). G+C content of its DNA is 37·0 mol%. Isolated from a bottom sediment sample collected in the Gulf of Peter the Great, Sea of Japan.

Description of Maribacter orientalis sp. nov.
Maribacter orientalis (o.ri.en.ta'lis. L. adj. orientalis eastern, bacterium inhabiting the East).

Main characteristics are the same as those given for the genus. In addition, cells range in size from 0·5 to 0·7 µm in width and 2 to 10 µm in length. On marine agar, colonies are 2–4 mm in diameter, circular, shiny with entire edges and yellow–orange pigmented. Grows at 4–32 °C; optimal temperature for growth is between 21 and 23 °C. Grows in the presence of 1–5 % NaCl. Decomposes gelatin, alginate, Tween 20 and Tween 40. Does not hydrolyse agar, casein, DNA, starch, urea, Tween 80, cellulose (carboximethylcellulose and filter paper) or chitin. Forms acid from arabinose, cellobiose, galactose, glucose, lactose, maltose, melibiose, sucrose and xylose, but not from fucose, raffinose, rhamnose, citrate, adonitol, dulcitol, glycerol, inositol or mannitol. Utilizes arabinose, glucose, lactose, mannose and sucrose, but not mannitol, inositol, sorbitol, malonate or citrate. Nitrate is not reduced. H2S, indole and acetoin (Voges–Proskauer reaction) production are negative. Susceptible to carbenicillin, lincomycin and oleandomycin; resistant to ampicillin, bensylpenicillin, gentamicin, kanamycin, neomycin, polymyxin B, streptomycin and tetracycline.

Type strain is KMM 3947T (=KCTC 12967T=CCUG 48008T). G+C content of its DNA is 39·0 mol%. Isolated from a sea water sample collected in the Gulf of Peter the Great, Sea of Japan.

Description of Maribacter aquivivus sp. nov.
Maribacter aquivivus (a.qui.vi'vus. L. fem. n. aqua water; N.L. adj. vivus alive; N.L. aquivivus living in water).

Main characteristics are the same as those given for the genus. In addition, cells range in size from 0·4 to 0·5 µm in width and 1·2 to 1·4 µm in length. On marine agar, colonies are 2–4 mm in diameter, circular, shiny with entire edges, yellow pigmented and sunken into agar. Grows at 4–30 °C; optimal temperature for growth is between 21 and 23 °C. Grows in the presence of 1–7 % NaCl. Decomposes agar, starch, alginate, Tween 20, Tween 40 and Tween 80. Does not hydrolyse gelatin, casein, DNA, cellulose (carboximethylcellulose and filter paper) or chitin. Does not form acid from arabinose, cellobiose, fucose, galactose, glucose, lactose, maltose, melibiose, raffinose, rhamnose, sucrose, xylose, citrate, adonitol, dulcitol, glycerol, inositol or mannitol. Utilizes glucose, lactose, mannose and sucrose, but not arabinose, mannitol, inositol, sorbitol, malonate or citrate. Nitrate is reduced. H2S, indole and acetoin (Voges–Proskauer reaction) production are negative. Susceptible to carbenicillin, lincomycin and oleandomycin; resistant to ampicillin, bensylpenicillin, gentamicin, kanamycin, neomycin, polymyxin B, streptomycin and tetracycline.

Type strain is KMM 3949T (=KCTC 12968T=CCUG 48009T). G+C content of its DNA is 35·0 mol%. Isolated from a sea water sample collected in the Gulf of Peter the Great, Sea of Japan.

Description of Maribacter ulvicola sp. nov.
Maribacter ulvicola (ul.vi.co'la. N.L. fem. n. Ulva generic name of green alga Ulva fenestrata; L. suffix -cola dweller; N.L. n. ulvicola a green alga Ulva fenestrata dweller).

Main characteristics are the same as those given for the genus. In addition, cells are flexible rods ranging in size from 0·25 to 0·3 µm in width and 4 to 6 µm in length. On marine agar, colonies are 2–4 mm in diameter, circular, shiny with entire edges, yellow pigmented and weakly sunken into agar. Grows at 4–32 °C; optimal temperature for growth is between 21 and 23 °C. Grows in the presence of 1–4 % NaCl. Does not decompose casein, gelatin, DNA, starch, alginate, urea, Tween 20, Tween 40, Tween 80, cellulose (carboximethylcellulose and filter paper) or chitin. Agar is hydrolysed weakly. Forms acid from cellobiose, fucose, glucose, lactose, maltose, rhamnose and sucrose, but not from arabinose, galactose, melibiose, raffinose, xylose, citrate, adonitol, dulcitol, glycerol, inositol or mannitol. Utilizes glucose, lactose, mannose and sucrose, but not arabinose, mannitol, inositol, sorbitol, malonate or citrate. Nitrate is not reduced. H2S, indole and acetoin (Voges–Proskauer reaction) production are negative. Susceptible to ampicillin, carbenicillin, lincomycin and oleandomycin; resistant to bensylpenicillin, gentamicin, kanamycin, neomycin, polymyxin B, streptomycin and tetracycline.

Type strain is KMM 3951T (=KCTC 12969T=DSM 15366T). G+C content of its DNA is 35–37 mol%. Isolated from the green alga Ulva fenestrata.


   ACKNOWLEDGEMENTS
 
We thank Professor Dr Hans G. Trüper and Dr Jean P. Euzéby for their advice on the species epithets. This research was supported by grants from the Ministry for Industry, Science and Technologies of the Russian Federation (no. 03-19) and Russian Foundation for Basic Research (no. 02-04-49517). K. S. B., S. K. H. and S. B. K. are also grateful for the support from the Korea Research Council of Fundamental Science and Technology (grant no. KBM1000212).


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Akagawa-Matsushita, M., Itoh, T., Katayama, Y., Kuraishi, H. & Yamasato, K. (1992). Isoprenoid quinone composition of some marine Alteromonas, Marinomonas, Deleya, Pseudomonas and Shewanella species. J Gen Microbiol 138, 2275–2281.

Barbeyron, T., L'Haridon, S., Corre, E., Kloareg, B. & Potin, P. (2001). Zobellia galactanovorans gen. nov., sp. nov., a marine species of Flavobacteriaceae isolated from red alga, and classification of [Cytophaga] uliginosa (ZoBell and Upham 1944Go) Reichenbach 1989Go as Zobellia uliginosa gen. nov., comb. nov. Int J Syst Evol Microbiol 51, 985–997.[Abstract]

Bowman, J. P. (2000). Description of Cellulophaga algicola sp. nov., isolated from the surfaces of Antarctic algae, and reclassification of Cytophaga uliginosa (ZoBell and Upham 1944Go) Reichenbach 1989Go as Cellulophaga uliginosa comb. nov. Int J Syst Evol Microbiol 50, 1861–1868.

Bowman, J. P. & Nichols, D. S. (2002). Aequorivita gen. nov., a member of the family Flavobacteriaceae isolated from terrestrial and marine Antarctic habitats. Int J Syst Evol Microbiol 52, 1533–1541.[Abstract]

Bruns, A., Rohde, M. & Berthe-Corti, L. (2001). Muricauda ruestringensis gen. nov., sp. nov., a facultatively anaerobic, appendaged bacterium from German North Sea intertidal sediment. Int J Syst Evol Microbiol 51, 1997–2006.[Abstract]

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Felsenstein, J. (1993). PHYLIP (phylogeny inference package), version 3.5c. Department of Genetics, University of Washington, Seattle, USA.

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