Subdivision of the genus Gluconacetobacter Yamada, Hoshino and Ishikawa 1998: the proposal of Komagatabacter gen. nov.

, for strains accommodated to the Gluconacetobacter xylinus group in the #-Proteobacteria Yuzo Yamada, Pattaraporn Yukphan, Huong Thi Lan Vu, Yuki Muramatsu, Duangjai Ochaikul & Yasuyoshi Nakagawa
Annals of Microbiology ISSN 1590-4261 Ann Microbiol DOI 10.1007/ s13213-011-0288-4

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and the type species of the new genus is designated as Komagatabacter xylinus (Brown 1886) comb. nov. Japan Y Yamada . Thailand Y Yamada . Kisarazu 292–0818.1007/s13213-011-0288-4 SHORT COMMUNICA TION Subdivision of the genus Gluconacetobacter Y amada. nov. . Bangkok 10520. 2-5-8 Kazusa Kamatari. Ward 4.ne. Shibuya-ku. Gluconacetobacter diazotrophicus (Gillis et al. an Hochiminh City. 1998. 1998.Author's personal copy Ann Microbiol DOI 10. King Mongkut’s Institute of Technology Ladkrabang. Vietnam National University–HCMC. Japan nacetobacter xylinus group. Y amada and Kondo (1985) did not consider that the two species were similar to . Acetobacter (Gluconacetobacter) liquefaciens (Asai 1935) Gosselé et al.jp H. 1992) Yamada et al. with the following five species. . Keywords Gluconacetobacter . T. 227 Nguyen V Cu Street. Gluconacetobacter liquefaciens (Asai 1935) Yamada et al. 1983 (the type species) and Acetobacter (Gluconacetobacter) xylinus (Brown 1886) Yamada 1984 were accommodated to the subgenus (Yamada and Kondo 1985). nov. 1998 (Yamada et al. phenotypically and ecologically: the Gluconacetobacter liquefaciens group and the GlucoY Yamada (*) : P Y . 113 Thailand Science Park. Klong 1. Klong Luang. Twelve species of the Gluconacetobacter xylinus group are transferred to the new genus as new combinations. Acetic acid bacteria . Vietnam . Acetobacteraceae Introduction The genus Gluconacetobacter Yamada et al. Gluconacetobacter xylinus (Brown 1886) Yamada et al. Komagatabacter gen. only two species. Ladkrabang. Hoshino and Ishikawa 1998: the proposal of Komagatabacter gen. Tokyo 151–8558. Y Muramatsu : Y Nakagawa . Upon the proposal of the subgenus Gluconacetobacter. NITE Biological Resource Center. 1998 and Gluconacetobacter europaeus (Sievers et al. However. Thailand e-mail: yamada333@kch. Phaholyothin Road. Shizuoka University. Suruga-ku. Faculty of Biology. 1997. Pathumthani 12120. Shizuoka 422–8529. Japan D. ukphan BIOTEC Culture Collection. National Science and Technology Development Agency. National Institute of Technology and Evaluation. Professor Emeritus. Japan International Cooperation Agency. Ochaikul Department of Biology. Vu Department of Microbiology. L. National Center for Genetic Engineering and Biotechnology. 1998). Faculty of Science. nov. Komagatabacter xylinus comb. University of Sciences. . 1983) Yamada et al. 1998 (the type species). Chalongkrung Road. 1989) Y amada et al. District 5. Gluconacetobacter hansenii (Gosselé et al.biglobe. the genus Komagatabacter is newly introduced. 1998 (Gluconoacetobacter sic) was introduced by an elevation of the subgenus Gluconacetobacter (ex Asai 1935) Yamada and Kondo 1985. JICA Senior Overseas V olunteer.. For the latter group. for strains accommodated to the Gluconacetobacter xylinus group in the α-Proteobacteria Yuzo Y amada & Pattaraporn Yukphan & Huong Thi Lan Vu & Yuki Muramatsu & Duangjai Ochaikul & Y asuyoshi Nakagawa Received: 8 February 2011 / Accepted: 20 May 2011 # Springer-V erlag and the University of Milan 2011 Abstract The genus Gluconacetobacter is divided into two groups phylogenetically.

in which G. and the other of G. oboediens. strains of this group were isolated mostly from fermented foods such as vinegar. 1998) Yamada 2000 and Gluconacetobacter intermedius (Boesch et al. 2000. In constructing a phylogenetic tree by the maximum likelihood method (Felsenstein 1981). 1983. This indicated that the two groups were phylogenetically ‘not so tightly coupled. 2005 belonged phylogenetically to the genus Gluconacetobacter. for example. Materials and methods All the 16S rRNA gene sequences used in this study are present in the GenBank/EMBL/DDBJ databases. Gluconobacter and Acetobacter. Leifson 1954. Distance matrices for the aligned sequences were calculated by the two-parameter method of Kimura (1980). nov. Chemotaxonomically. G. namely the Gluconacetobacter xylinus group and the Gluconacetobacter liquefaciens group. Yamada and Y ukphan (2008) suggested that the Gluconactobacter liquefaciens group and the Gluconacetobacter xylinus group in the genus Gluconacetobacter can be distinguished from each other at the generic level from the phylogenetical. as found in strains of species of the genus Gluconobacter.6. in which G. Neoasaia. Lisdiyanti et al. Yamada et al.Author's personal copy Ann Microbiol each other chemotaxonomically and phenotypically (Asai et al. Phenotypically. Phylogenetic trees based on 16S rRNA gene sequences of 1. europaeus had been located. the program PHYLIP (version 3. 2007). in contrast to those of 70. Gluconacetobacter sacchari Franke et al. J. (2000) calculated the 16S rRNA gene sequence similarities between the type strains of species belonging to Gluconacetobacter subclusters 1 and 2 to be 96. 74 and 60% in the genus Gluconacetobacter. G. 2001 and Gluconacetobacter johannae Fuentes-Ramírez et al. fruits. 1999. liquefaciens. Multiple alignments of the cited DNA sequences were performed with the program Clustal X (version 1. intermedius. This paper proposes Komagatabacter gen. coffee plants and so on. in addition to the above-mentioned isolation sources. Kozakia. 2006. Alignment gaps and unidentified bases were eliminated. 2005 and Gluconacetobacter rhaeticus Dellaglio et al. (2006) described that the species of the genus Gluconacetobacter seemed likely to be divided into two subclusters phylogenetically.5-diketo.D -gluconate. The calculated bootstrap value was 60%. swingsii. 2006. namely. Y amada (2000) transferred Acetobacter oboediens and Acetobacter intermedius to Gluconacetobacter subcluster 1. G. diazotrophicus.. but the latter showed only Q-10. a major component of Q-9(Q-8) in strains of species of the genus Acetobacter.0. Tamura et al.8. 1964. Thompson et al. Gluconacetobacter saccharivorans Lisdiyanti et al. Yamada 1976. nata de coco. in the Gluconacetobacter xylinus branch. the family Acetobacteraceae Gillis and De Ley 1980.5–97. but the Gluconacetobacter xylinus group is not necessarily plant-associated.3%. using the program MEGA (version 4. rhaeticus. Gluconacetobacter nataicola Lisdiyanti et al. 2001.’ (2) The Gluconacetobacter liquefaciens group formed a large cluster along with the genera Acidomonas. (5) Ecologically. University of Washington) was . europaeus. in which Q-9. as Gluconacetobacter oboediens (Sokollek et al. strains of the Gluconacetobacter liquefaciens group produced 2. For example. hansenii and G. the calculated bootstrap values were 100% in the neighborjoining method. hansenii and Gluconacetobacter entanii Schüller et al. a minor component of the respiratory quinone homologues. 1997). the former was motile with peritrichous flagella. namely strains of this group were isolated mostly from flowers. liquefaciens and G. G. Asaia. 1998) Yamada 2000. G. 91% in the maximum parsimony method and 90% in the maximum likelihood method.219 bases were constructed by the neighbor-joining method (Saitou and Nei 1987) and the maximum parsimony method (Felsenstein 1983). G. on the basis of results obtained by 16S rRNA gene sequence analyses and by phenotypic characterization. (3) A similar clustering was found in a 16S rRNA gene sequence phylogenetic tree derived from the maximum likelihood method. Such a clustering was never found in acetic acid bacteria. but strains of the Gluconacetobacter xylinus group did not. tea fungus beverages and so on.6% between the two subclusters. xylinus. but not to the Gluconacetobacter subcluster 2. the Gluconacetobacter liquefaciens group is plant-associated. for strains of the species accommodated to the Gluconacetobacter xylinus group of the genus Gluconacetobacter. diazotrophicus had been located. Gluconacetobacter azotocaptans Fuentes-Ramírez et al. The species accommodated to the genus Gluconacetobacter are divided into two groups as mentioned above. Swaminathania. xylinus. (4) Physiologically. the former presented Q-10(Q-9). 1969). Yamada et al. one consisting of G.7–97. corresponded to Q-9. G. rather than the cluster of the Gluconacetobacter xylinus group in a 16S rRNA gene sequence phylogenetic tree derived from the maximum parsimony method. The calculated pair-wise 16S rRNA gene sequence similarities were 95. sugarcane. Dellaglio et al. (2005) reported that the two isolates that classified as Gluconacetobacter swingsii Dellaglio et al. γ-pyrone compounds and a water-soluble brown pigment. phenotypical and ecological points of view: (1) The calculated bootstrap value between the two groups in a 16S rRNA gene sequence phylogenetic tree derived from the neighbor-joining method was 70%. The calculated bootstrap value was 74%. G. in the genus Gluconobacter. Felsenstein. but the latter was non-motile.

as indicated previously (Yamada and Y ukphan 2008). which contained 2. and bootstrap values are not shown if below 50% in phylogenetic trees. 2006). and that produce biocellulose. 2000) and Y ukphan et al. 60 and 71% in the phylogenetic trees derived from the neighbour-joining method. the former was plant-associated. (1964). One included Acetobacter aceti (the type species). (2002. the 12 species constituted a cluster in which species that strictly require acetic acid for growth. 2008. the calculated bootstrap value was 66%. and the other included Acetobacter pasteurianus. Yamada et al. (1983). 1). 1–3). The results obtained differed from those of Dutta and Gachhui (2007). as suggested by Y amada and Y ukphan (2008). 2001). the MP method and the ML method.0% agar w/v. As described above. europaeus and G. The type strain of G.000 replications (Felsenstein 1985). but the latter was not necessarily plant-associated. These phylogenetic.2% yeast extract w/v. the five species classified in the Gluconacetobacter liquefaciens group were divided into two subclusters. azotocaptans. the Gluconacetobacter liquefaciens group and the Gluconacetobacter xylinus group were differentiated phylogenetically from each other at the generic level. (2005. Physiologically. Between the two clusters. 0. Between the clusters of the Glconacetobacter liquefaciens group and the Glconacetobacter xylinus group. Acetobacter peroxydans and so on. nataicola. Dutta and Gachhui (2007) reported that the type strain of G. were likely to be distributed randomly. The robustness of individual branches was estimated by bootstrapping with 1. The type strain of Acidocella facilis was used as an outgroup. However. In the Gluconacetobacter xylinus group. Lisdiyanti et al.5-diketo-D-gluconate and γ-pyrone compounds from D-glucose. Furthermore. the species assigned to the Gluconacetobacter xylinus group were phylogenetically independent of the Gluconacetobacter liquefaciens group. the type strain of the species was examined for motility and flagellation. 1. (1969. Ecologically.0). groEL and rpoB. but the latter did not. Motility was tested by the hanging-drop method. G. The bacterial strain was grown at 20°C for 18–20 h on an agar plate. 2). phenotypic and ecological characteristics are enough to separate the Gluconacetobacter xylinus group from the Gluconacetobacter liquefaciens group at the generic level (Table 1) (Yamada and Y ukphan 2008). Acetobacter estunensis and so on.0% glucose w/v. The two subclusters were connected to each other with a bootstrap value of 97%. such as G. and there did not appear to be any rule governing their distribution (Table 1). The former was not characterized by a nitrogen fixation capability. 2011). but the latter was (Gillis et al. 0. the former produced a water soluble-brown pigment when grown on glucose/yeast extract/calcium carbonate medium while the latter did not. johannae and G. they stated that some of the above-mentioned phenotypic features were of little use for the differentiation of acetic .Author's personal copy Ann Microbiol used instead of the program MEGA (version 4. In all the three phylogenetic trees based on 16S rRNA gene sequences. Phenotypic features were determined by the methods of Asai et al.5% peptone w/v. diazotrophicus. (2010) described that the genus Gluconacetobacter should not remain as a single genus on the basis of multilocus sequence analyses of the three housekeeping genes of dnaK. Cleenwerck et al. sacchari and the other including the three species. kombuchae assigned to the Gluconacetobacter xylinus group. the former was motile with peritrichous flagella. the calculated bootstrap values were 72. G. The cluster of the Gluconacetobacter xylinus group showed similar clustering. The type strain of Gluconacetobacter kombuchae of the Gluconacetobacter xylinus group was examined for morphology. and cells were negatively stained and observed under an electron microscope. such as G. liquefaciens and G. and no flagellation was found (data not shown). a similar clustering was found in both the Gluconacetobacter liquefaciens group and the Gluconacetobacter xylinus group. kombuchae (= LMG 23726T) was nonmotile. The morphological property of the type strain observed in this study was reasonable and consistent with those of strains of the species assigned to the Gluconacetobacter xylinus group (Table 1). 2009. G. entanii. Navarro and Komagata (1999). kombuchae was motile with polar flagellation. as found in the phylogenetic tree constructed by the neighbor-joining method. It was noted that the 19 species assigned to the genus Acetobacter represented a similar clustering to the genus Gluconacetobacter (Figs. Biochemically. one comprising the two species.7% calcium carbonate w/v and 2. 3).0% ethanol v/v. and the latter was non-motile without any flagellation. Considering the morphology of G. In a phylogenetic tree based on 16S rRNA gene sequences derived from the maximum parsimony (MP) method (Fig. 0. The calculated bootstrap value was 42% between the two clusters. xylinus and G. 1976. Gosselé et al. the five species classified in the Gluconacetobacter liquefaciens group also grouped into two subclusters or subgroups. In a phylogenetic tree based on 16S rRNA gene sequences derived from the maximum likelihood (ML) method (Fig. respectively. The calculated bootstrap value between the two clusters was 53%. Morphologically. 1989. the former produced 2. Results and discussion In a phylogenetic tree based on 16S rRNA gene sequences derived from the neighbor-joining method (Fig. Fuentes-Ramírez et al.

however. one can . K. Our opinion is. quite different from theirs. The type strain of Acidocella facilis was used as an outgroup.Author's personal copy Ann Microbiol Fig. Komagatabacter acid bacteria and that. other differentiating features should be looked for before splitting the genus. In fact.000 replications. for this reason. The above-mentioned phenotypic features are useful for differentiating the Gluconacetobacter xylinus group from the Gluconacetobacter liquefaciens group. Numerals at nodes indicate bootstrap values (%) derived from 1. A phylogenetic tree based on 16S rRNA gene sequences was constructed by the neighbor-joining method. 1 Phylogenetic relationships of the genera Gluconacetobacter and Komagatabacter.

7 + + + + + + + + + + + + + + + + + + + + − + nd + + nd + − − − + nd + + nd + nd nd + − nd + − nd nd + + − 63. no none. 2005).8g + + − 59. 1999). nataicola LMG 1536T (Lisdiyanti et al. − negative. intermedius strain TF2T (Boesch et al. 2006).Ann Microbiol Table 1 Differential characteristics of the genera Gluconacetobacter and Komagatabacter. TH TH europaeus strain DES 11T (Sievers et al. 7 K. 2001). + positive.4 per per per per no no no no no no no no 2 3 4 5 6 7 8 9 10 11 12 13 14 no + + + + + + − + − − + − + − − 61 Komagatabacter 15 no + + + + + + − + + − + − + + − 62 16 nob + + + nd nd + − nd + − nd nd − + − 55. Gluconacetobacter strains: 1 Gluconacetobacter liquefaciens NBRC 12388T (Navarro and Komagata 1999). rhaeticus strain DST GLO2T (Dellaglio et al. 2001). johannae strain CFN-Cf55T (Fuentes-Ramírez et al. 2006).3 nd nd + − − 59.8 17 no + + + + + + − + + − nd nd + − − 62. 2000). 1995. Lisdiyanti et al. 1998).9 + + + − +b + + − − +b + + nd − +b + + nd − +b + − + + − + − + − − + − nd − − + − + − − + − nd − − nd nd − − − 61. saccharivorans LMG 1582T (Lisdiyanti et al. 2005). 1998). 2000 f Yamada et al. hansenii NBRC 14820T (Lisdiyanti et al. entanii strain L 4560T (Schüller et al. kombuchae strain RG3T (Dutta and Gachhui 2007). 2006). 1981 g Tanaka et al. 14 K. 4 G. azotocaptans strain CFNCa54T (Fuentes-Ramírez et al. a major ubiquinone was Q-10 in all strains tested b Present study c Navarro and Komagata 1999. 2010) Gluconacetobacter 1 per + + + + + + + + − + + −b. 2000 . 3 G. 11 K. 2 G. 1992).5-Diketo-D-gluconate DNA G+C content (mol%) a per peritrchous. Cleenwerck et al. swingsii strain DST GL01T (Dellaglio et al. 17 K.6 + + + + nd nd nd nd + + + + + + nd nd + + nd nd + − nd − − nd nd − − − 58 nd nd + − nd + − nd nd + + − 61. 1989). 2006 d Lisdiyanti et al. 16 K. Komagatabacter strains: 6 Komagatabacter xylinus JCM 7644T (Navarro and Komagata 1999). 15 K. 12 K. 9 K. sacchari strain SRI 1794T (Franke et al.0 + + − 61. 5 G.0 + + + + − + nd nd + − − + nd nd + + − 56.0 +b +b +b 64. 8 K. sucrofermentans strain BPR 2001T (Toyosaki et al.e + + + 64.5f + − + 61 + + + 65 +b +b +b 58. 2006 e Lisdiyanti et al. diazotrophicus LMG 7603T (Gillis et al. oboediens L 2460T (Sokollek et al. nd not determined.2-57.3 Characteristica Flagellation Oxidation of: Acetate Lactate Growth without acetic acid Growth on:c Glutamate agar Mannitol agar Author's personal copy Production of acetic acid from ethanol Production of a water-soluble brown pigment Production of dihydroxyacetone from glycerol Cellulose production Production of γ-pyrone compound Assimilation of ammoniac nitrogen on:d Mannitol Ethanol Production of: 2-Keto-D-gluconate 5-Keto-D-gluconate 2. 10 K. 13 K.

retention index=0.Author's personal copy Ann Microbiol Fig. when it shows the oxidation of acetate and lactate and produces a characteristic water-soluble brown pigment (Table 1). The isolate . Komagatabacter exactly classify or identify such an acetic acid bacterium. K. of which 129 were parsimony informative. Numerals at nodes indicate bootstrap values (%) derived from 1. A phylogenetic tree based on 16S rRNA gene sequences was constructed by the maximum parsimony (MP) method.795. specifically to the Gluconacetobacter liquefaciens group but not to the Gluconacetobacter xylinus group. There were a total of 1. as has already been done by Lisdiyanti et al. 2 Phylogenetic relationships of the genera Gluconacetobacter and Komagatabacter. The type strain of Acidocella facilis was used as an outgroup. by the use of the phenotypic features mentioned above. (2004) and Greenberg et al. Loganathan and Nair (2004). without any problem and without any confusion. (2002).351. homoplasy index= 0.000 replications. Jojima et al.441. For example. Consistency index=0. The phylogenetic relationships were represented by a consensus tree of two most parsimonious trees. (2006) (see Y amada and Y ukphan 2008).559. a certain isolate can be assigned to or classified into the genus Gluconacetobacter. rescaled consistency index=0.219 positions (bases) in the final dataset.

A phylogenetic tree based on 16S rRNA gene sequences was constructed by the maximum likelihood (ML) method. in general.Author's personal copy Ann Microbiol Fig. all of which have no oxidation capability of acetate and lactate. . 2011). and also from strains of the genus Swaminathania that have weak acetate and lactate oxidation. although all of them produce the water-soluble brown pigment (Y ukphan et al. 3 Phylogenetic relationships of the genera Gluconacetobacter and Komagatabacter. In contrast. The type strain of Acidocella facilis was used as an outgroup. K.000 replications. Komagatabacter also can be differentiated from some strains of the genus Gluconobacter and strains of the genus Tanticharoenia. an isolate that is non-motile and does not produce the water-soluble brown pigment can. Numerals at nodes indicate bootstrap values (%) derived from 1.

maltose or lactose. Production of dihydroxyacetone from glycerol is positive or negative. 64.5. Since the new genus was introduced. especially of acetic acid bacteria). which is characterized by little production of acetic acid from ethanol. L. grows on glutamate agar and mannitol agar. but not from D-fructose. the acetate and lactate oxidation test is useful (Asai et al. Production of cellulose is positive or negative. which is characterized by Q-9 as a major quinone homologue. The former produces a water-soluble brown pigment. and the species of the former group can appropriately be classified into a separate new genus. but the latter does not. Gluconobacter or Asaia. acetic acid is strictly required for growth. differing from the genus Gluconacetobacter . in isolates to be assigned to the genus Acetobacter. Y amada and Y ukphan 2008). Japan. neut. The name of the genus is Komagatabacter gen. In some strains. the characteristic brown-pigment production can be utilized. nov. N. since there are likely to be no decisive phenotypic features to discriminate the two clusters or the two groups taxonomically. Gluconacetobacter. D-fructose or D-mannitol. 2008. measuring 0. nov. Grows on D-glucose. 1964.bac’ter. the systematic assignments or the taxonomic positions of the isolate based on routine identification methods can be confirmed phylogenetically. only five genera of acetic acid bacteria include more than one species.0–3. Kozakia and Granulibacter that show weak acetate oxidation. the color change to blue will be less vigorous compared to isolates to be assigned to the genus Acetobacter. 2011). A major isoprenoid quinone is Q-10. but not on lactose. In general. In some strains. namely Acetobacter Beijerinck 1898. Gluconobacter Asai 1935. Y ukphan et al. are isolated from the natural environments.ta. Kazuo Komagata. acetate and lactate are not oxidized. 2009. n. At the present time. color change will be very slow. Tokyo.5-diketo-D-gluconate is not produced. which are oxidative but not fermentative and grow at pH 3. however. Int J Syst Bacterial 48: .8–63. Produces 2-keto-D-gluconate or 5-keto-D-gluconate from D-glucose.8 × 1. Hoshino and Ishikawa. nov. the following species should be transferred to the genus Komagatabacter. Others are monotypic genera and seem likely to be composed of rare microorganisms in distribution. D-galactose.Author's personal copy Ann Microbiol be assigned to or classified into the Gluconacetobacter xylinus group of the genus Gluconacetobacter. In isolates to be assigned to the genus Gluconacetobacter. from strains of the genera Asaia. Isolates to be assigned to the genus Gluconobacter will generally show a clear yellow color. In some strains. Thus. To discriminate these acetic acid bacteria at the generic level. To divide isolates already assigned to the genus Gluconacetobacter into two groups. N. Komagatabacter (Ko. DNA base composition is 55. a deep blue color will appear fast and clearly.5– 0. nov. from strains of the genus Acidomonas that grow on methanol by showing no growth on methanol. The isolate also can be differentiated from strains of the genus Acetobacter that have Q-9 and the quite intense ability to oxidize acetate and lactate to carbon dioxide and water. namely the Gluconacetobacter liquefaciens group and the Gluconacetobacter xylinus group. Biosci Biotechnol Biochem 61: 1250. bacter from Gr. γ-Pyrone compounds are not produced. D-xylose. Neoasaia. Gram-negative rods and non-motile. In the case of isolates to be assigned to the genus Asaia. The University of Tokyo. L-sorbose. Oxidizes acetate and lactate to carbon dioxide and water. The type species is Komagatabacter xylinus (Brown 1886) comb. who contributed to the bacterial systematics. To date.ga. masc. but 2. Does not produce a water-soluble brown pigment on glucose/yeast extract/calcium carbonate medium. It is noteworthy that. masc. The 19 species assigned to the genus Acetobacter to date represent two clusters with low bootstrap values at the branching points in the phylogenetic trees (Figs. baktron rod. n. L-arabinose or ethanol.4 mol%G+C with a range of 7. Description of Komagatabacter gen. ketogluconates are not produced.6 mol%.1–3) (Y amada and Y ukphan 2008). The phylogenetic and phenotypic characteristics obtained above are enough to separate the Gluconacetobacter xylinus group at the generic level from the Gluconacetobacter liquefaciens group. Asaia Yamada et al.35% acetic acid v/v. For example. Synonym: Gluconacetobacter xylinus (Brown 1886) Y amada. n. Saccharibacter and Neokomagataea that have no oxidation of acetate and lactate (Yamada and Y ukphan 2008. Komagatabacter xylinus (Brown 1886) comb. When acetic acid bacteria. for example. derived from Dr. 2011. from strains of the genus Ameyamaea that are motile with polar flagella and weak oxidation of lactate and from the genera Gluconobacter. the species of the two clusters cannot be differentiated at the generic level. Finally.ma. when it shows oxidation of acetate and lactate and has Q-10 as a major isoprenoid quinone (Table 1). in our experience. Acid is produced from D-glucose. one can identify a large number of isolates in a short period. 2000 and Neokomagataea Y ukphan et al. Produces acetic acid from ethanol. Komagatabacter Komagata rod. there have been no exceptions to this routine identification method. one will encounter mostly acetic acid bacteria that are classified into either the genus Acetobacter . Ammoniac nitrogen is generally assimilated on D-mannitol. D-sorbitol. D-mannitol. Growth is positive in the presence of 0. L. by constructing a phylogenetic tree based on 16S rRNA gene sequences. 1997 (V alidation list no.0 μm. Bunkyo-ku. Colonies are white–creamy and smooth with entire margin or rough. Professor Emeritus.

Pauwels and De Ley. Synonym: Guconacetobacter hansenii Gosselé. nov. Tsuchida. Int J Syst Evol Microbiol 59:1771–1786 Cleenwerck I. Komagatabacter intermedius (Boesch. Kojima. J Gen Appl Microbiol 41:312. Uchimura and Komagata. sucrofermentans Toyosaki. Int J Syst Bacteriol 30: 239. Int J. Sellmer and Teuber 1992) comb. Synonym: Gluconacetobacter sucrofermentans (Toyosaki. Felis. Hoshino and Ishikawa. Trček. 1992 (V alidaton list no. Trček. Y amada and Y oshinaga 1996) Cleenwerck. Felis. Komagatabacter swingsii (Dellaglio. According to Lisdiyanti et al. Komagatabacter europaeus (Sievers. 2007. Sellmer and Teuber. Sievers and Teuber 1998) Y amada. Acetobacter xylinus (Brown 1886) Y amada. The type strain is NCIMB 8746T. The type strain is LMG 23726T. 12. Komagatabacter oboediens (Sokollek. Navarro. Komagatabacter nataicola (Lisdiyanti. Komagatabacter rhaeticus (Dellaglio. 2005. Int J Syst Bacterial 33: 896. Komagatabacter entanii (Schüller. 64. Hertel and Hammes. Basonym: Gluconacetobacter rhaeticus Dellaglio. Tsuchida. 64. Navarro. Engelbeen. Hoshino and Ishikawa. nov. Basonym: Gluconacetobacter swingsii Dellaglio. Int J Syst bacterial 42: 656. 67. Engelbeen. Pauwels and De Ley 1983) Y amada. Swings. Int J Syst Evol Microbiol 60: os 2282. this species is a later heterotypic synonym of Gluconacetobacter oboediens. Janssens and Marzotto. nov. Synonym: Gluconacetobacter intermedius (Boesch. The type strain is LMG 22125T. (2006). 1998) Basonym: Acetobacter hansenii Gosselé. Felis. Navarro. Hoshino. of species of the genus Gluconacetobacter and related taxa based . 2010 Basonym: Acetobacter xylinus (Brown 1886) Yamada 1984 subsp. nov. nov. Cleenwerck. 1992). this species is a later heterotypic synonym of Gluconacetobacter hansenii. Janssens and Marzotto 2005) comb. Engelbeen. Syst Appl Microbiol 4: 366. Komagatabacter saccharivorans (Lisdiyanti. Cleenwerck. Kojima. References Asai T. 1998. The type strain is LMG 1536T. Int J Syst Bacterial 48: 939. De V P De Vuyst L (2010) Phylogeny and differentiation os . The type strain is DSM 13536T. Trček J. Hoshino. 2006. Pauwels and De Ley 1983) comb. 1995 (V alidation list no. Komagata K (1964) The flagellation and taxonomy of genera Gluconobacter and Acetobacter with reference to the existence of intermediate strains. The type strain is DSM 11804T. Syst Evol Microbiol 50: 226. Sievers and Teuber 1998) comb. 1984) Basonym: Acetobacter aceti subsp. Int J Syst Bacteriol 48: 327. De Vuyst L. 1996). Int J Syst Evol Microbiol 50: 2019. Basonym: Gluconacetobacter entanii Schüller Hertel and Hammes. Y amada and Yoshinaga. Iizuka H. Int J Syst Bacteriol 46: 836. Hertel and Hammes 1998) comb. Teuber M (1998) Acetobacter intermedius sp. 58. Sievers M. De V P os (2009) Differentiation of species of the family Acetobacteraceae by AFLP DNA fingerprinting: Gluconacetobacter kombuchae is a later heterotypic synonym of Gluconacetobacter hansenii. Syst Evol Microbiol 50: 226. (2009). Int J Syst Bacteriol 34: 270. J Gen Appl Microbiol 29: 419. Kersters. nov. Swings. xylinus (xylinum sic) (Brown 1886) De Ley and Frateur 1974 (Approved lists. Trček. J Gen Appl Microbiol 10:95–126 Boesch C. According to Cleenwerck et al. 1998). The type strain is LMG 1582T. The type strain is NCIMB 11664T. Kersters. Cleenwerck. Int J Syst Evol Microbiol 55: 2369. Y amada and Y oshinaga 1996) comb. De Wachter M. Syst Appl Microbiol 15: 391. Basonym: Gluconacetobacter kombuchae Dutta and Gachhui. 2000 Basonym: Acetobacter intermedius Boesch. 2000. Biosci Biotechnol Biochem 61: 1250. Basonym: Gluconacetobacter nataicola Lisdiyanti. 2000 Basonym: Acetobacter oboediens Sokollek. Janssens and Marzotto 2005) comb. Hertel and Hammes 1998) Y amada. The type strain is JCM 9730T. Basonym: Gluconacetobacter saccharivorans Lisdiyanti. Hoshino. nov. 1983 (V alidation list no. Int J. González Á. 2006. Sellmer and Teuber 1992) Y amada. 14. 1997 (V alidation list no. Tsuchida. Cleenwerck. 1998 (V alidation list no. Swings. Komagatabacter kombuchae (Dutta and Gachhui 2007) comb. Int J Syst Evol Microbiol 57: 356. Sievers and Teuber. Uchimura and Komagata 2006) comb. Kersters. Komagatabacter hansenii (Gosselé. The type strain is DSM 6160T. The type strain is DSM 11826T. Synonym: Gluconacetobacter europaeus (Sievers. Uchimura and Komagata 2006) comb. Janssens and Marzotto. 1998). 1980). Synonym: Gluconacetobacter oboediens (Sokollek. Syst Appl Microbiol 21: 228. 1983). nov. Int J Syst Evol Microbiol 55: 2368. Int J Syst Bacterial 48: 327. Int J Syst Bacteriol 48: 1083. The type strain is LMG 22126T. 1983 (V alidation list no. Int J Syst Evol Microbiol 56: 2108. 2005. Biosci Biotechnol Biochem 61: 1250.Author's personal copy Ann Microbiol 327. Komagatabacter sucrofermentans (Toyosaki. nov. Kojima. Hertel and Hammes 2000) comb. Felis. 1997 (V alidation list no. nov. V and Vuyst. nov. Syst Appl Microbiol 21:220–229 Cleenwerck I. 43. Uchimura and Komagata. Engelbeen. nov. 1998) Basonym: Acetobacter europaeus Sievers. Int J Syst Evol Microbiol 56: 2109. Navarro.

a salt-tolerant. nov. Conville PS. Yamashita M. Gibson TJ. J Gen Appl Microbiol 22:237–245 Yamada Y Ishikawa T. Janssens D. Aoki K (2000) Genetic characteristics of cellulose-forming acetic acid bacteria identified phenotypically as Gluconacetobacter xylinus. a novel acetic acid bacterium in the α-Proteobacteria. J Gen Appl Microbiol 22:285–292 Y amada Y (1983) Acetobacter xylinus sp.. Tapia-Hernández A. Janssens D. J Mol Evol 17:368–376 Felsenstein J (1983) Parsimony in systematics: biological and statistical issues. Porcella SF. Int J Syst Evol Microbiol 55:2365–2370 Dutta D.. Int J Food Microbiol 125:15–24 Yamada Y Aida K. Int J Syst Bacteriol 49:1681–1693 Fuentes-Ramírez LE. nov. sp. nov.. nov. a nitrogen-fixing acetic acid bacterium associated with sugarcane. a new subgenus . Yamada Y Yoshinaga F . and Acetobacter estunensis (Carr 1958) comb. nov. 1998 to the genus Gluconacetobacter as Gluconacetobacter oboediens comb. Hammes WP (2000) Gluconacetobacter entanii sp. ukphan P (2008) Genera and species in acetic acid bacteria. sp. nov. Nei M. Martínez-Romero E. nov. Jeanmougin F. nov. Bustillos-Cristales R. Y . J Gen Appl Microbiol 30:297–303 Yamada Y Y . Fudou R . a new species of acetic acid bacterium isolated from the leaf sheath of sugar cane and from the pink sugar-cane mealy bug.. Leonard G. Stephan MP Teixeira KRS. Dudley J. sp. nom. Döbereiner J. two new species isolated from industrial vinegar fermentations. nov. sp. Zelazny AM (2006) Granulibacter bethesdensis gen. Yamada Y . J Mol Evol 16:111–120 Leifson E (1954) The flagellation and taxonomy of species of Acetobacter. CaballeroMellado J (2001) Novel nitrogen-fixing acetic acid bacteria. Kaneko T . “polarly flagellated intermediate strains” in acetic acid bacteria. Int J Syst Evol Microbiol 54:1185–1190 Navarro RR. Plewniak F. Mol Biol Evol 24:1596–1599 Tanaka M. Fegan M. Int J Syst Evol Microbiol 54:2263–2267 Kimura M (1980) A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. nov. J Gen Appl Microbiol 27:465–475 . nov. Tahara Y Yamasato K. a distinctive pathogenic acetic acid bacterium in the family Acetobacteraceae. analysis of phenotypic features and protein gel electrophoregrams of a wide variety of Acetobacter strains. nov. Marzotto M (2005) Description of Gluconacetobacter swingsii sp. acetate-oxidizing acetic acid bacteria with the Q-10 system. Gluconacetobacter johannae sp. for the cellulose-forming and cellulose-less. Felis GE. 1998 and Acetobacter intermedius Boesch et al... Evolution 39:783–791 Franke IH. Hammes WP (1998) Description of Acetobacter oboediens sp. amada Y Uchimura T. . . Gachhui R (2007) Nitrogen-fixing and cellulose-producing Gluconacetobacter kombuchae sp.. Murray PR. Stackebrandt E.. J Gen Appl Microbiol 29:417–420 Yamada Y (2000) Transfer of Acetobacter oboediens Sokollek et al.Author's personal copy Ann Microbiol on multilocus sequence analyses of housekeeping genes and reclassification of Acetobacter xylinus subsp. nov. associated with coffee plants. especially in the polarly flagellated intermediate strains. Nucleic Acids Res 25:4876–4882 Toyosaki H. Shinke R.. sification of Gluconacetobacter hansenii strains and proposals of Gluconacetobacter saccharivorans sp. Nei M (1987) The neighbor-joining method: a new method for reconstructing phylogenetic trees. . Komagata K .. Teuber M (1992) Acetobacter europaeus sp.. Uchimura T. nov. and Gluconacetobacter rhaeticus sp. Holland SM. Komagata K (2006) Reclas. Kersters K. nov. nov. Int J Syst Evol Microbiol 56:2609– 2616 Jojima Y Mihara Y Suzuki S. . Uemura T (1969) Enzymatic studies on the . Kumar S (2007) MEGA4: Molecular evolutionary genetics analysis (MEGA) software version 4... nov. J Gen Appl Microbiol 41:307–314 Yamada Y (1976) Characterization of Acetobacter xylinum by ubiquinone system. (1981) Deoxyribonucleic acid base composition and deoxyribonucleic acid homology in acetic acid bacteria. Syst Appl Microbiol 15:386–392 Sokollek SJ. nov. Annu Rev Ecol Syst 14:313–333 Felsenstein J (1985) Confidence limits on phylogenies: an approach using the bootstrap.. Hayward C. Antonie V Leeuwenhoek 20:102–110 an Lisdiyanti P Kawasaki H. 215. V Ubiquinone of acetic . Kroppenstedt RM. nov. . Int J Syst Evol Microbiol 50:2225–2227 Y amada Y Kondo K (1985) Gluconoacetobacter. isolated from Kombucha tea. Seki T. nov. Hertel C. Acetobacter tropicalis sp. Seki T. Int J Syst Evol Microbiol 57:353–357 Felsenstein J (1981) Evolutionary trees from DNA sequences: a maximum likelihood approach. Hoshino K.. Hertel C. nov. nov.. Hoste B. 1996) sp. isolated from submerged high-acid industrial vinegar fermentations. nov.. J Gen Appl Microbiol 15:186–196 Yamada Y Okada Y Kondo K (1976) Isolation and characterization of . Kojima Y Tsuchida T. Kersters K. Int J Syst Evol Microbiol 51:1305–1314 Gillis M. Komagata K (1999) Differentiation of Gluconacetobcter liquefaciens and Gluconacetobacter xylinus on the basis of DNA base composition. Murakami S. nov. Int J Syst Evol Microbiol 56:2101–2111 Loganathan P Nair S (2004) Swaminathania salitolerans gen..0. Proposal for the improvement of the taxonomy of the genus Acetobacter Beijerinck 1898. (2004) Saccharibacter floricola gen. Stock F. nov. a novel osmophilic acetic acid bacterium isolated from pollen. Sly LI (1999) Description of Gluconacetobacter sacchari sp. Int J Syst Evol Microbiol 52:813–818 Lisdiyanti P Navarro RR. Wang ET. . nov. and Acetobacter pomorum sp. nov.. nov. Jiménes-Salgado T. Sellmer S. Yokozeki K. sucroferementans as Gluconacetobacter sucrofermentans (Toyosaki et al. comprising the acetate-oxidizing acetic acid bacteria with ubiquinone-10 in the genus Acetobacter. Pauwels P De Ley J (1983) Numerical . Swings J. Komagata K (2002) Kozakia baliensis gen. Acetobacter orleanensis (Henneberg 1906) comb. Int J Syst Bacteriol 48:935–940 Tamura K. Mol Biol Evol 4:406–425 Schüller G. De Ley J (1989) Acetobacter . nov. Engelbeen K. Syst Appl Microbiol 4:338–368 Greenberg DE. Widyastuti Y Saono S. Int J Syst Evol Microbiol 60:2277–2283 Dellaglio F. (1995) The characterization of an acetic acid bacterium useful for producing bacterial cellulose in agitation cultures: the proposal of Acetobacter xylinum subsp. (2000) Systematic study of the genus Acetobacter with descriptions of Acetobacter indonesiensis sp. and Gluconacetobacter azotocaptans sp. nitrogen-fixing and phosphate-solubilizing bacterium from wild rice (Porteresia coarctata Tateoka). Cleenwerck I. nov. Biosci Biotechnol Biochem 64:757–760 Thompson JD. nov.. Acetobacter lovaniensis (Frateur 1950) comb. nov. Int J Syst Evol Microbiol 50:2013–2020 Sievers M. Wong A. J Gen Appl Microbiol 45:7–15 Saitou N.. isolated from Italian apple fruit. sucrofermentans subsp. oxidation of sugar and sugar alcohol. DNA relatedness and oxidation products from glucose. and Gluconacetobacter intermedius comb. diazotrophicus sp. Uchimura T. Higgins DG (1997) The CLUSTAL X windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. and Gluconacetobacter nataicola sp. especially of the so-called intermediate strains. Yamanaka S.. J Gen Appl Microbiol 46:147–165 Lisdiyanti P Kawasaki H. Int J Syst Bacteriol 39:361–364 Gosselé F. . rev. acid bacteria and its relation to classification of genera Gluconobacter and Acetobacter. comb. a main component of industrial vinegar fermenters in central Europe.

. nov.. Ishikawa T (1998) V . Uchimura T. Kaneyasu M. Y . Nakagawa Y Hamana K. Potacharoen W. alidation list no.. nov. sp. Tanticharoen M. Yamada . Int J Syst Evol Microbiol 50:823–829 Y ukphan P Malimas T. osmotolerant acetic acid bacteria of the α-Proteobacteria. Tanasupawat S. sp. Muramatsu Y Takahashi M. a new osmotolerant acetic acid bacterium in the α-Proteobacteria. with descriptions of Neokomagataea thailandica sp. Komagata K (2000) Asaia bogorensis gen. Potacharoen W. an acetic acid bacterium in the α-Proteobacteria. Muramatsu Y Takahashi M. nov. 64. Tanasupawat S. Kaneyasu M. .. sp. . acid bacteria based on the partial sequences of 16S ribosomal RNA: the elevation of the subgenus Gluconoacetobacter to the generic level. nov. Biosci Biotechnol Biochem 73:2156–2162 Y ukphan P Malimas T. nov.. Y amada Y (2005) Neoasaia chiangmaiensis gen. Tanticharoen M. Tahara . Biosci Biotechnol Biochem 72:672–676 Y ukphan P Malimas T. Yamada Y (2009) Ameyamaea . . chiangmaiensis gen. amada Y (2011) Neokomagataea gen. Tanasupawat S.. Y (2008) Tanticharoenia sakaeratensis gen.. Biosci Biotechnol Biochem 75:419–426 .Author's personal copy Ann Microbiol Yamada Y Hoshino K. . . . Nakagawa Y Suzuki K. Int J Syst Bacteriol 48:327– 328 Yamada Y Katsura K. nov. an unusual acetic acid bacterium in the α-Proteobacteria. a novel osmotolerant acetic acid bacterium in the α-Proteobacteria. Y Suzuki K. nov. nov. .. nov.. Nakagawa Y Tanticharoen M. Kawasaki H. J Gen Appl Microbiol 51:301–311 Y ukphan P Malimas T. and Neokomagataea tanensis sp.. Potacharoen W. . Widyastuti Y Saono S. nov. sp. Biosci Biotechnol Biochem 61:1244–1251 Y amada Y Hoshino K. nov. Muramatsu Y Potacharoen W Tanasupawat S. Seki T. V alidation of publication of new names and new combinations previously effectively published outside the IJSB. Ishikawa T (1997) The phylogeny of acetic .. .

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