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Int J Syst Evol Microbiol 54 (2004), 157-162; DOI  10.1099/ijs.0.02861-0
© 2004 International Union of Microbiological Societies

Hongiella mannitolivorans gen. nov., sp. nov., Hongiella halophila sp. nov. and Hongiella ornithinivorans sp. nov., isolated from tidal flat sediment

Hana Yi and Jongsik Chun

School of Biological Sciences, Seoul National University, 56-1 Shillim-dong, Kwanak-gu, Seoul 151-742, Republic of Korea

Correspondence
Jongsik Chun
chun{at}snu.ac.kr


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Three marine strains of the Cytophaga–Flavobacterium–Bacteroides group, designated JC2050T, JC2051T and JC2052T, were obtained from a single sediment sample of getbol, the Korean tidal flat. Comparative 16S rDNA sequence studies revealed that the test strains were not closely related to any validly published genera and that these strains were only distantly related to the genus Cyclobacterium (88·7–91·2 %). Phylogenetic analyses demonstrated that the three getbol isolates formed a distinct monophyletic clade within the family Cytophagaceae. Physiological, biochemical and chemotaxonomic data also indicated that these three getbol isolates differed significantly from members of other genera and were sufficiently different from each other to be recognized as separate species. On the basis of polyphasic evidence, a new genus, Hongiella gen. nov., is proposed, with three novel species, Hongiella mannitolivorans sp. nov. (type strain JC2050T=IMSNU 14012T=DSM 15301T), Hongiella halophila sp. nov. (type strain JC2051T=IMSNU 14013T=DSM 15292T) and Hongiella ornithinivorans sp. nov. (type strain JC2052T=IMSNU 14014T=DSM 15282T). Hongiella mannitolivorans is the type species of the genus.

Abbreviations: CFB, Cytophaga–Flavobacterium–Bacteroides

Published online ahead of print on 18 July 2003 as DOI 10.1099/ijs.0.02861-0.

The GenBank accession numbers for 16S rDNA sequences of strains JC2050T, JC2051T and JC2052T are respectively AY264838, AY264839 and AY264840.


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
During a course of study on the culturable aerobic bacterial community in getbol (Korean tidal flat), a large number of novel bacterial strains was isolated (Yi & Chun, 2002Go; Yi et al., 2003Go). Three of these isolates were members of the Cytophaga–Flavobacterium–Bacteroides (CFB) lineage and were the subject of a taxonomic investigation. On the basis of polyphasic evidence, these test strains formed a new genus in the CFB group, for which the name Hongiella gen. nov. is proposed.


   METHODS
TOP
 ABSTRACT
 INTRODUCTION
 METHODS
RESULTS AND DISCUSSION
 REFERENCES
 
Bacterial strains.
Test strains were isolated from a single sample of tidal flat sediment from Ganghwa Island, Korea (37°35'31·9''N; 126°27'24·5''E). Strains JC2050T and JC2052T were isolated on marine agar 2216 (MA; Difco) and strain JC2051T was isolated on MR2A [R2A (Difco) supplemented with artificial sea salts (Sigma)]. The isolates were routinely cultured on MA at 30 °C and maintained as a glycerol suspension (20 %, w/v) at -80 °C.

Molecular systematics.
Bacterial DNA preparation, PCR amplification and 16S rDNA sequencing were carried out as described previously (Chun & Goodfellow, 1995Go). The resultant sequences were aligned manually against sequences obtained from GenBank. Phylogenetic trees were inferred using the Fitch–Margoliash (Fitch & Margoliash, 1967Go), maximum-likelihood (Felsenstein, 1993Go), maximum-parsimony (Fitch, 1972Go) and neighbour-joining (Saitou & Nei, 1987Go) methods. Evolutionary distance matrices were generated according to Jukes & Cantor (1969)Go. The resultant tree topologies were evaluated in bootstrap analyses (Felsenstein, 1985Go) based on 1000 resamplings. Alignment and phylogenetic analyses were carried out using the programs PHYDIT (available at http://plaza.snu.ac.kr/~jchun/phydit/) and PAUP 4.0 (Swofford, 1998Go) as described previously (Chun et al., 2000Go).

Cultural, morphological and physiological properties.
Cultural characteristics were studied using several bacteriological growth media: CY (3 g casitone, 1 g yeast extract, 1 g CaCl2.2H2O, 40 g sea salts and 15 g agar in 1000 ml distilled water), CSY-3 (Sawabe et al., 1998Go); 1/5 LBM (Suzuki et al., 2001Go), MA, SMM (Shioi's marine medium; Shiba, 1992Go) and YTSS (Gonzalez et al., 1997Go). Growth at various temperatures was tested over the range 5–50 °C at intervals of 5 °C using MA. Growth at different pH values (4–14) was examined on MA. Growth in NaCl (0–12 % at 0·5 % intervals) or sea salts (0–15 % at 0·5 % intervals) was tested on CSY-3.

Cellular morphology was examined by phase-contrast microscopy and SEM after growth on MA at 30 °C. Gliding motility was observed by direct microscopic examination of the edge of colonies and motility was observed by the hanging-drop technique for cells in exponential phase in CY broth.

Carbon-source utilization was tested on 96-well tissue-culture microplates as described by Gosink et al. (1998)Go. Strain JC2051T was able to grow on Baumann's basal medium (BM; Baumann et al., 1971Go) without added growth factors, but strains JC2050T and JC2052T were unable to grow on BM without trace elements and vitamins. Thus, BM, supplemented with 2 % (v/v) Hutner's mineral base (Cohen-Bazire et al., 1957Go) and 1 % (v/v) vitamin solution (Staley, 1968Go), was used for carbon-source testing. {beta}-Galactosidase activity was determined by streaking cultures onto MA agar plates supplemented with 0·1 mM IPTG and 20 µg X-Gal ml-1 (Gosink et al., 1998Go). Other physiological and biochemical properties were tested using standard procedures as described previously (Yi et al., 2003Go).

Chemotaxonomy.
Chemotaxonomic characteristics were determined in cells grown at 30 °C for 2 days on MA or in MB (marine broth 2216; Difco). The presence of flexirubin-like pigments was tested by measuring the absorbance spectrum of an ethanol and alkaline-ethanol extract of lysed cells (Weeks, 1981Go). Fatty acid methyl ester analysis was performed by GLC according to the instructions of the Microbial Identification system (MIDI). Isoprenoid quinones were isolated according to Minnikin et al. (1984)Go and analysed using HPLC as described by Collins (1985)Go. G+C content (mol%) was determined by HPLC analysis of deoxyribonucleosides as described by Mesbah et al. (1989)Go using a reverse-phase column (Supelcosil LC-18-S; Supelco).


   RESULTS AND DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 METHODS
 RESULTS AND DISCUSSION
REFERENCES
 
Phylogenetic analysis
Nearly complete 16S rDNA sequences of strains JC2050T (1376 bp), JC2051T (1427 bp) and JC2052T (1425 bp) were obtained and used for an initial BLAST search against GenBank. The result clearly indicated that the getbol isolates belonged to the family Cytophagaceae within the CFB group. The newly determined sequences were then aligned manually against representatives of the CFB group based on secondary structure of bacterial 16S rRNA. Sequence similarities among the three isolates were 94–96 %, all below the cut–off value of 97 % for bacterial species definition (Stackebrandt & Goebel, 1994Go). The highest similarity values obtained between our isolates and species with validly published names were observed with Cyclobacterium marinum ATCC 43824 (88·7 % for JC2050T, 90·1 % for JC2051T and 91·2 % for JC2052T). No other validly named bacterial species showed more than 90 % 16S rDNA sequence similarity to the test strains. The distinctiveness of our isolates was also evident in the phylogenetic tree (Fig. 1Go). Strains JC2050T and JC2052T formed a monophyletic clade with 94 % bootstrap support. Strain JC2051T formed a sister group to this clade, and all three isolates formed a monophyletic clade supported by a bootstrap value of 100 %. Cyclobacterium marinum ATCC 43824 was recovered as a further sister group with a bootstrap value of 100 %. Tree topology was supported by all tree-making methods used in this study. However, the branching patterns of the other genera varied depending on the tree-building method used and the relationships were supported by relatively low bootstrap values. On the basis of low 16S rDNA similarity values (<92 %) between our isolates and known genera and their significantly monophyletic grouping, it is fair to conclude that the test strains should be placed in a new genus of the CFB group.



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  		Fig. 1. Phylogenetic relationship of strains JC2050T, JC2051T and JC2052T and related taxa within the CFB group based on 16S rDNA sequences. The tree was created using the neighbour-joining method and numbers at nodes are levels of bootstrap support (%) from 1000 resampled datasets. Solid circles indicate that the corresponding nodes (groupings) are also recovered in Fitch–Margoliash, maximum-likelihood and maximum-parsimony trees. Helicobacter pylori ATCC 43504T (GenBank accession number U01330) was used as an outgroup (not shown). Bar, 0·1 nt substitution per position.

 
Culture and growth conditions
Abundant growth was observed on CSY-3, MA and SMM. When grown on CY, 1/5 LBM and YTSS agar, smaller colonies with smaller amounts of pigment were observed. Strains grew at temperatures of 10–40 °C (optimum 35–40 °C) for JC2050T, 5–40 °C (optimum 35 °C) for JC2051T and 10–45 °C (optimum 35–40 °C) for JC2052T. Although all of the test strains were isolated from a marine habitat, only strain JC2051T showed slight halophilicity, as it grew on media containing 1–10 % (w/v) NaCl (optimum 5 % NaCl) or 0·5–11 % (w/v) artificial sea salts-based media (optimum 1–2 % sea salts). Strains JC2050T and JC2052T were able to grow without NaCl or sea salts; strain JC2050T grew at 0–7 % NaCl (optimum 1 % NaCl) or 0–7 % sea salts (optimum 0·5–1·5 % sea salts) and strain JC2052T grew at 0–10 % NaCl (optimum 1 % NaCl) or 0–13 % sea salts (optimum 1·0–2·5 % sea salts). Optimum growth occurred at pH 7; the pH range for growth was pH 6–11 for strain JC2050T and pH 6–13 for strains JC2051T and JC2052T. No growth was detected under anaerobic conditions generated by a GasPack system (BBL).

Morphological properties
Colonies of strains JC2050T and JC2052T were circular, convex, glistening, non-luminescent, butyraceous, opaque and orange with entire margins on agar plates. Colonies of strain JC2051T were slightly different, i.e. flat, translucent and reddish-orange. Cells were rod-shaped with rounded ends (Fig. 2Go), non-flagellated and non-motile. Gliding motility was not observed on agar plates, nor was motility observed in broth media. Spore formation was not observed.



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  		Fig. 2. SEM of strain JC2050T. Cells were grown on MA at 30 °C for 2 days. Bar, 1 µm.

 
Physiological and biochemical properties
The physiological and biochemical properties of the test strains are given in Table 1Go. A summary of the major differences between the test strains and phylogenetically related species is given in Table 3Go.


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  		Table 1. Phenotypic features of strains JC2050T, JC2051T and JC2052T

+, Positive; -, negative; W, weakly positive. All three strains were positive for catalase, oxidase, hydrolysis of aesculin and X-Gal, alkaline phosphatase, leucine arylamidase, acid phosphatase, naphthol-AS-BI-phosphohydrolase, {alpha}-galactosidase and N-acetyl-{beta}-glucosaminidase. All three strains were negative for arginine dihydrolase, fluorescein, fermentation of glucose, polyhydroxybutyrate accumulation, urease, production of acetoin, H2S and indole, hydrolysis of agar, alginic acid, casein, cellulose, chitin and egg yolk, esterase (C4), esterase lipase (C8), lipase (C14), cystine arylamidase, {alpha}-mannosidase and {alpha}-fucosidase. The following compounds were utilized as sole carbon sources by all test strains: D-cellobiose, D-glucose, D-mannose, D-raffinose, D-salicin, D-trehalose, D-xylose, lactose, N-acetylglucosamine and sucrose. The following compounds were not utilized by any of the three strains: acetamide, acetate, benzoate, citrate, D-ribose, D-sorbitol, ethanol, glycerol, glycine, inositol, inulin, 2-propanol, L-arabinose, L-arginine, L-ascorbate, L-asparagine, L-lysine, polyethylene glycol, salicylate, succinate, tartrate and thiamin.

 

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  		Table 3. Characteristics that can be used to differentiate strains JC2050T, JC2051T and JC2052T from phylogenetically related species

Genera: 1, Hongiella (strains JC2050T, JC2051T and JC2052T); 2, Cyclobacterium; 3, Cytophaga; 4, Dyadobacter; 5, Flectobacillus; 6, Flexibacter; 7, Runella; 8, Spirosoma; 9, Sporocytophaga. Data are from this study and others (Chelius & Triplett, 2000Go; Fujita et al., 1996Go; Gosink et al., 1998Go; Raj & Maloy, 1990Go; Reichenbach, 1989Go; Urakami & Komagata, 1986Go). +, Positive; -, negative; W, weakly positive; V, variable; ND, no data available.

 
Chemotaxonomic characteristics
Absorption maxima of crude extracts of strains JC2050T and JC2052T were approximately 480 nm; that of strain JC2051T was 475 nm. Bathychromatic shift, a characteristic of flexirubins, was not observed in any strains. The predominant cellular fatty acids of the three strains were 15 : 0 iso, iso 17 : 1{omega}9c, 17 : 0 iso 3-OH and mixture of 16 : 1{omega}7c and 15 : 0 iso 2-OH (Table 2Go). The overall fatty acid profiles of our isolates differed significantly from those in phylogenetically related taxa, except for the genus Runella (Table 3Go). However, the latter showed a much higher DNA G+C content (49–50 mol%) than our isolates (37–42 mol%). MK-7 was the predominant isoprenoid quinone in all test strains. The DNA G+C contents of strains JC2050T, JC2051T and JC2052T were respectively 42, 37 and 38 mol%.


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  		Table 2. Cellular fatty acid content of strains JC2050T, JC2051T and JC2052T

Values are percentages of total fatty acid content. Only fatty acids representing at least 1 % of the total fatty acids of at least one of the strains are shown.

 
Taxonomic conclusions
Polyphasic analyses demonstrated that the three getbol isolates belong to a coherent group that corresponds to a novel genus within the CFB group. Phenotypic data clearly demonstrated that test strains are not closely affiliated with any previously described genera (Table 3Go) and are sufficiently different from each other to be recognized as separate species (Table 1Go). In many aspects, strain JC2051T was shown to be rather distantly related to the other two strains, namely colonial morphology, NaCl requirement, growth capability on BM and the maximum absorption of pigment. The 16S rDNA sequence similarity and the phylogenetic tree also reflected this relationship.

Based on the polyphasic data presented in this study, the name Hongiella gen. nov. is proposed for the getbol isolates, with three novel species, Hongiella mannitolivorans sp. nov. for strain JC2050T, Hongiella halophila sp. nov. for strain JC2051T and Hongiella ornithinivorans sp. nov for strain JC2052T.

Description of Hongiella gen. nov.
Hongiella (Hong.i.el'la. N.L. dim. fem. n. Hongiella named after Soon-Woo Hong, a Korean microbiologist who devoted his life to the study of soil micro-organisms).

Gram-negative, oxidase- and catalase-positive. Strictly aerobic, chemoheterotrophic and mesophilic. Grows optimally at neutral pH. Cells are rod-shaped with rounded ends, non-flagellated and non-motile. Colonies are circular, entirely margined, glistening, butyraceous and orange on MA and do not glide. Abundant growth occurs on CSY-3, MA and SMM media. Spores are not formed. Flexirubin-type pigment is absent. The major isoprenoid quinone is MK-7. The predominant cellular fatty acids are 15 : 0 iso (23–29 %), iso 17 : 1{omega}9c (6–9 %), 17 : 0 iso 3-OH (8–11 %) and a mixture of 16 : 1{omega}7c and 15 : 0 iso 2-OH (14–21 %). DNA G+C content is 37–42 mol%. Many phenotypic characters differentiate this genus from related taxa (Table 3Go). Phylogenetically, the genus belongs to the family Cytophagaceae. The type species is Hongiella mannitolivorans.

Description of Hongiella mannitolivorans sp. nov.
Hongiella mannitolivorans (man.ni.to.li.vo'rans. N.L. n. mannitolum mannitol; L. v. vorare to devour; N.L. part. adj. mannitolivorans utilizing mannitol).

Cells are approximately 1·1–1·7x0·4–0·5 µm. Colonies are convex, opaque and orange on MA. Optimal growth is observed at 35–40 °C, pH 7 and 1 % NaCl or 0·5–1·5 % artificial sea salts. Grows without NaCl or sea salts. Reduces nitrate to nitrite. Produces amylase, DNase and gelatinase, but not Tweenase. Detailed physiological and biochemical characteristics are given in Table 1Go. Maximum absorption of pigment occurs at 480 nm. Cellular fatty acid composition is given in Table 2Go. The type strain is JC2050T (=IMSNU 14012T=KCTC 12050T=DSM 15301T); the DNA G+C content of the type strain is 42 mol%. Isolated from sediment of getbol, the Korean tidal flat.

Description of Hongiella halophila sp. nov.
Hongiella halophila (ha.lo'phi.la. Gr. n. hals, halos salt; Gr. adj. philos loving; N.L. fem. adj. halophila salt-loving).

Cells are approximately 1·0–1·8x0·3–0·5 µm. Colonies are flat, translucent and reddish-orange on MA. Optimal growth is observed at 35 °C, pH 7 and 5 % NaCl or 1–2 % artificial sea salts. Does not grow without NaCl or sea salts. Nitrate is not reduced to nitrite. Produces gelatinase and Tweenase, but not amylase or DNase. Detailed physiological and biochemical characteristics are given in Table 1Go. Maximum absorption of pigment occurs at 475 nm. Cellular fatty acid composition is given in Table 2Go. The type strain is JC2051T (=IMSNU 14013T=KCTC 12051T=DSM 15292T); the DNA G+C content of the type strain is 37 mol%. Isolated from sediment of getbol, the Korean tidal flat.

Description of Hongiella ornithinivorans sp. nov.
Hongiella ornithinivorans (or'ni.thi.ni.vo'rans. N.L. n. ornithinum ornithine; L. v. vorare to devour; N.L. part. adj. ornithinivorans utilizing ornithine).

Cells are approximately 0·8–2·6x0·3–0·4 µm. Colonies are convex, opaque and orange on MA. Optimal growth is observed at 35–40 °C, pH 7 and 1 % NaCl or 1·0–2·5 % artificial sea salts. Grows without NaCl or sea salts. Nitrate is not reduced to nitrite. Produces amylase, DNase, gelatinase (weakly) and Tweenase. Detailed physiological and biochemical characteristics are given in Table 1Go. Maximum absorption of pigment occurs at 480 nm. Cellular fatty acid composition is given in Table 2Go. The type strain is JC2052T (=IMSNU 14014T=KCTC 12052T=DSM 15282T); the DNA G+C content of the type strain is 38 mol%. Isolated from sediment of getbol, the Korean tidal flat.


   ACKNOWLEDGEMENTS
 
We are grateful to Dr J. P. Euzéby for help with nomenclature. This work was supported by the 21C Frontier Microbial Genomics and Applications Center Program (grant MG02-0101-001-2-1-0) and the Strategic National R&D Program through the Genetic Resources and Information Network Center, MOST (grant M1-0219-00-0018). H. Y. was supported by a BK21 Research Fellowship.


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 Int. J. Syst. Evol. Microbiol.Home page
Q.-M. Liu, L. N. Ten, H.-S. Yu, F.-X. Jin, W.-T. Im, and S.-T. Lee
Emticicia ginsengisoli sp. nov., a species of the family 'Flexibacteraceae' isolated from soil of a ginseng field
Int J Syst Evol Microbiol, May 1, 2008; 58(5): 1100 - 1105.
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 Int. J. Syst. Evol. Microbiol.Home page
J. L. Copa-Patino, M. Arenas, J. Soliveri, C. Sanchez-Porro, and A. Ventosa
Algoriphagus hitonicola sp. nov., isolated from an athalassohaline lagoon
Int J Syst Evol Microbiol, February 1, 2008; 58(2): 424 - 428.
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 Int. J. Syst. Evol. Microbiol.Home page
O. I. Nedashkovskaya, S. B. Kim, K. K. Kwon, D. S. Shin, X. Luo, S.-J. Kim, and V. V. Mikhailov
Proposal of Algoriphagus vanfongensis sp. nov., transfer of members of the genera Hongiella Yi and Chun 2004 emend. Nedashkovskaya et al. 2004 and Chimaereicella Tiago et al. 2006 to the genus Algoriphagus, and emended description of the genus Algoriphagus Bowman et al. 2003 emend. Nedashkovskaya et al. 2004
Int J Syst Evol Microbiol, September 1, 2007; 57(9): 1988 - 1994.
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Y. Zhou, X. Wang, H. Liu, K.-Y. Zhang, Y.-Q. Zhang, R. Lai, and W.-J. Li
Pontibacter akesuensis sp. nov., isolated from a desert soil in China
Int J Syst Evol Microbiol, February 1, 2007; 57(2): 321 - 325.
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H.-W. Chang, Y.-D. Nam, H.-Y. Kwon, J. R. Park, J.-S. Lee, J.-H. Yoon, K.-G. An, and J.-W. Bae
Marinobacterium halophilum sp. nov., a marine bacterium isolated from the Yellow Sea
Int J Syst Evol Microbiol, January 1, 2007; 57(1): 77 - 80.
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K. Suresh, S. Mayilraj, and T. Chakrabarti
Effluviibacter roseus gen. nov., sp. nov., isolated from muddy water, belonging to the family 'Flexibacteraceae'
Int J Syst Evol Microbiol, July 1, 2006; 56(Pt 7): 1703 - 1707.
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O. I. Nedashkovskaya, S. B. Kim, M. Vancanneyt, A. M. Lysenko, D. S. Shin, M. S. Park, K. H. Lee, W. J. Jung, N. I. Kalinovskaya, V. V. Mikhailov, et al.
Echinicola pacifica gen. nov., sp. nov., a novel flexibacterium isolated from the sea urchin Strongylocentrotus intermedius.
Int J Syst Evol Microbiol, May 1, 2006; 56(Pt 5): 953 - 958.
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J.-H. Yoon, M.-H. Lee, S.-J. Kang, and T.-K. Oh
Algoriphagus terrigena sp. nov., isolated from soil.
Int J Syst Evol Microbiol, April 1, 2006; 56(Pt 4): 777 - 780.
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 Int. J. Syst. Evol. Microbiol.Home page
L. A. O'Sullivan, J. Rinna, G. Humphreys, A. J. Weightman, and J. C. Fry
Culturable phylogenetic diversity of the phylum 'Bacteroidetes' from river epilithon and coastal water and description of novel members of the family Flavobacteriaceae: Epilithonimonas tenax gen. nov., sp. nov. and Persicivirga xylanidelens gen. nov., sp. nov.
Int J Syst Evol Microbiol, January 1, 2006; 56(1): 169 - 180.
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H.-Y. Weon, B.-Y. Kim, S.-W. Kwon, I.-C. Park, I.-B. Cha, B. J. Tindall, E. Stackebrandt, H. G. Truper, and S.-J. Go
Leadbetterella byssophila gen. nov., sp. nov., isolated from cotton-waste composts for the cultivation of oyster mushroom
Int J Syst Evol Microbiol, November 1, 2005; 55(6): 2297 - 2302.
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O. I. Nedashkovskaya, S. B. Kim, M. Suzuki, L. S. Shevchenko, M. S. Lee, K. H. Lee, M. S. Park, G. M. Frolova, H. W. Oh, K. S. Bae, et al.
Pontibacter actiniarum gen. nov., sp. nov., a novel member of the phylum 'Bacteroidetes', and proposal of Reichenbachiella gen. nov. as a replacement for the illegitimate prokaryotic generic name Reichenbachia Nedashkovskaya et al. 2003
Int J Syst Evol Microbiol, November 1, 2005; 55(6): 2583 - 2588.
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J.-H. Yoon, S.-J. Kang, and T.-K. Oh
Algoriphagus locisalis sp. nov., isolated from a marine solar saltern
Int J Syst Evol Microbiol, July 1, 2005; 55(4): 1635 - 1639.
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A. H. Rickard, A. T. Stead, G. A. O'May, S. Lindsay, M. Banner, P. S. Handley, and P. Gilbert
Adhaeribacter aquaticus gen. nov., sp. nov., a Gram-negative isolate from a potable water biofilm
Int J Syst Evol Microbiol, March 1, 2005; 55(2): 821 - 829.
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O. I. Nedashkovskaya, S. B. Kim, D. H. Lee, A. M. Lysenko, L. S. Shevchenko, G. M. Frolova, V. V. Mikhailov, K. H. Lee, and K. S. Bae
Roseivirga ehrenbergii gen. nov., sp. nov., a novel marine bacterium of the phylum 'Bacteroidetes', isolated from the green alga Ulva fenestrata
Int J Syst Evol Microbiol, January 1, 2005; 55(1): 231 - 234.
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S. Van Trappen, I. Vandecandelaere, J. Mergaert, and J. Swings
Algoriphagus antarcticus sp. nov., a novel psychrophile from microbial mats in Antarctic lakes
Int J Syst Evol Microbiol, November 1, 2004; 54(6): 1969 - 1973.
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I. Brettar, R. Christen, and M. G. Hofle
Aquiflexum balticum gen. nov., sp. nov., a novel marine bacterium of the Cytophaga-Flavobacterium-Bacteroides group isolated from surface water of the central Baltic Sea
Int J Syst Evol Microbiol, November 1, 2004; 54(6): 2335 - 2341.
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O. I. Nedashkovskaya, M. Vancanneyt, S. Van Trappen, K. Vandemeulebroecke, A. M. Lysenko, M. Rohde, E. Falsen, G. M. Frolova, V. V. Mikhailov, and J. Swings
Description of Algoriphagus aquimarinus sp. nov., Algoriphagus chordae sp. nov. and Algoriphagus winogradskyi sp. nov., from sea water and algae, transfer of Hongiella halophila Yi and Chun 2004 to the genus Algoriphagus as Algoriphagus halophilus comb. nov. and emended descriptions of the genera Algoriphagus Bowman et al. 2003 and Hongiella Yi and Chun 2004
Int J Syst Evol Microbiol, September 1, 2004; 54(5): 1757 - 1764.
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J.-H. Yoon, S.-H. Yeo, and T.-K. Oh
Hongiella marincola sp. nov., isolated from sea water of the East Sea in Korea
Int J Syst Evol Microbiol, September 1, 2004; 54(5): 1845 - 1848.
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