International Journal of Systematic and Evolutionary Microbiology (2014), 64, 3473–3477 DOI 10.1099/ijs.0.066902-0

Tangfeifania diversioriginum gen. nov., sp. nov., a representative of the family Draconibacteriaceae

Qian-Qian Liu,1 Xiao-Li Li,1 Alejandro P. Rooney,2 Zong-Jun Du1,3 and Guan-Jun Chen1,3

Correspondence 1College of Marine Science, Shandong University at Weihai, Weihai 264209, PR China Zong-Jun Du 2National Center for Agricultural Utilization Research, Agricultural Research Service, [email protected] US Department of Agriculture, Peoria, IL 61604, USA 3State key Laboratory of Microbial Technology, Shandong University, Jinan 250100, PR China

A novel Gram-stain-negative, facultatively anaerobic, catalase- and oxidase-positive, non-motile and pink-pigmented bacterium, designated G22T, was isolated from Gahai, a saltwater lake in Qinghai province, China. Optimal growth occurred at 33–35 6C, pH 7.0–7.5, and in the presence of 2–4 % (w/v) NaCl. The DNA G+C content was 40.0 mol%. The major polar lipids were phosphatidylethanolamine and three unknown lipids. The predominant cellular fatty acids

were iso-C15 : 0, anteiso-C15 : 0, iso-C17 : 0 3-OH and iso-C15 : 0 3-OH, and MK-7 was the main respiratory quinone. Phylogenetic analysis based on 16S rRNA gene sequences revealed that strain G22T fell within the class Bacteroidia. Its closest phylogenetic neighbour was the recently described species Draconibacterium orientale, the sole member of the family Draconibacteriaceae, with merely 90.04 % sequence similarity. On the basis of phenotypic, chemotaxonomic and phylogenetic evidence observed, a novel species in a new genus, Tangfeifania diversioriginum gen. nov., sp. nov., is proposed within the family Draconibacteriaceae. The type strain is G22T (5CICC 10587T5DSM 27063T).

At the time of writing, the class Bacteroidia (phylum another novel family, Draconibacteriaceae, was created to ) comprises seven families: Bacteroidaceae, accommodate the newly described genus Draconibacterium , Porphyromonadaceae, Prevotellaceae, (Du et al., 2014). In this paper, we report the characterization Rikenellaceae, Prolixibacteraceae and Draconibacteriaceae. of a novel species in a new genus within the family Draconi- During the last decade, many novel taxa belonging to bacteriaceae, for which we propose the name Tangfeifania this class have been described, of which the phylogenetic diversioriginum gen. nov., sp. nov. placement for many is still unresolved, such as the genera Strain G22T was isolated from a sediment sample collected Marinifilum and Phocaeicola (Al Masalma et al., 2009; Na + 2 from Gahai, a saltwater lake (35 g Na l 1, 100u 329 330 E et al., 2009; Ruvira et al., 2013). By contrast, the classifica- 36u 599 500 N) located in Qinghai Province, China. The tion of certain taxa has been re-evaluated, resulting in a sediment sample was treated with an enrichment culture clarification of many outstanding problems. For instance, technique described by Du et al. (2014), except that the the misclassified [Cytophaga] fermentans has recently been plates were incubated at 28 uC for 2 weeks. Subsequently, a reclassified as fermentans within the family pink colony was isolated and purified through repeated Marinilabiliaceae (Yang et al., 2014). Likewise, the genera streaking on marine agar 2216 (MA; Difco) at 33 uC. Pure Prolixibacter and Sunxiuqinia (Holmes et al., 2007; Qu, cultures were stored at 280 uC in 20 % (v/v) glycerol or et al., 2011; Takai, et al., 2013), which represent deep lineages maintained as lyophilized pellets in vacuum-sealed glass in the phylum that were unable to be classified systematically vials stored at 4 uC. for many years, were re-evaluated and transferred to the family Prolixibacteraceae (Huang et al., 2014). Recently, Cell morphology was observed using light microscopy (E600; Nikon). Catalase activity was determined by observ- Abbreviations: ML, maximum-likelihood; MP, maximum-parsimony; NJ, ing bubble production in 3 % (v/v) H2O2 solution and neighbour-joining. oxidase activity was determined using an oxidase reagent The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene (bioMe´rieux). Gliding motility was examined by using oil- sequence of strain G22T is JQ683777. immersion phase-contrast microscopy (AX70; Olympus) as Three supplementary figures are available with the online Supple- described by Bowman (2000). The optimal growth tem- mentary Material. perature and pH and tolerance of NaCl were studied as

066902 Printed in Great Britain 3473 Q.-Q. Liu and others described previously (Du et al., 2014). Gram staining, unknown lipid as the major polar lipids, and MK-7 as the reduction of nitrate, the oxidation–fermentation test and main respiratory quinone. tests for degradation of agar, starch, CM-cellulose, alginate DNA was extracted and purified using a genomic DNA and Tween 80 were performed and evaluated by using MA extraction kit (Tiangen) following the manufacturer’s medium as described by Dong & Cai (2001). Additional protocol. The G+C content of DNA was determined by biochemical tests were carried out by using API 20E, API ZYM strips and API 50 CHB fermentation kits (bioMe´rieux) HPLC (Mesbah et al., 1989) and found to be 40.0 mol%. and Biolog GEN III microplates according to the manu- The 16S rRNA gene was amplified by PCR with universal facturers’ instructions, except that the NaCl concentration primers 27f and 1492r (Qu et al., 2011). The amplified gene was adjusted in all tests to 3 %. Antimicrobial susceptibility was cloned into pGM-T vector (Tiangen) and recombinant testing was performed by the disc diffusion method, as plasmids were reproduced in Escherichia coli DH5a cells. described by Jorgensen et al. (1999), using antibiotic- Sequencing reactions were carried out using an ABI BigDye impregnated discs (Tianhe) on MA incubated at 33 uC for 3.1 Sequencing kit (Applied BioSystems) and an automated 7 days. Results of these phenotypic tests are given in the DNA sequencer (model ABI3730; Applied BioSystems). species description and in Table 1. The resulting 16S rRNA gene sequence was submitted to GenBank and EMBL to search for similar sequences using Fatty acid compositions were determined as described the BLAST algorithm; in addition, the EzTaxon-e database by Sasser (1990) by using the Microbial Identification was queried in order to identify the most similar 16S rRNA System (Microbial ID). Cultures for fatty acid analysis were gene sequences from verified type strains (Kim et al., 2012). incubated in MB at 33 uC for 7 days. Isoprenoid quinones The sequence was aligned via CLUSTAL X (version 1.81) were extracted, purified and analysed by HPLC, as described (Thompson et al., 1997) with related sequences retrieved by Hiraishi et al. (1996). Polar lipids analysis was carried from the databases, and the alignments were edited manually. out by the Identification Service, Leibniz-Institut DSMZ – A phylogenetic tree was reconstructed using the neighbour- Deutsche Sammlung von Mikroorganismen und Zellkulturen, joining (NJ) method implemented in MEGA version 6 Braunschweig, Germany. (Tamura et al., 2013). The maximum-likelihood (ML) The predominant cellular fatty acids (.5 %) of strain G22T method, as implemented in MEGA 6, and the maximum- were iso-C15 : 0 (32.6 %), anteiso-C15 : 0 (15.4 %), iso-C17 : 0 parsimony (MP) method, as implemented in PAUP (Swofford, 3-OH (14.6 %) and iso-C15 : 0 3-OH (5.4 %), which was 2002), were also used to reconstruct phylogenetic trees in similar to those seen in Draconibacterium orientale except order to determine the phylogenetic placement of the novel T that G22 possessed more iso-C15 : 0 3-OH and less iso- isolate and reference species on the tree. The resultant tree C17 : 0 2-OH. topologies generated from all three methods were evaluated by bootstrap analysis based on 1000 replicates. The major polar lipids were phosphatidylethanolamine and three unknown lipids. In addition, moderate to minor The nearly complete 16S rRNA gene sequence (1446 nt) of amounts of glycolipid, aminolipid, two unknown phos- strain G22T was obtained. From the results of initial 16S pholipids and another unknown lipid were detected (Fig. rRNA gene sequence analysis by BLAST, some uncultured S1, available in the online Supplementary Material). The bacterial clones were found showing 93–99 % sequence main respiratory quinone was MK-7. By comparison, D. similarity to G22T. However, those studies (Alain et al., orientale also contains phosphatidylethanolamine and an 2006; Liu et al., 2009; Chang et al., 2012; Harris et al., 2013;

Table 1. Differential characteristics of strain G22T and related genera of the class Bacteroidia

Genera: 1, Tangfeifania gen. nov. (strain G22T); 2, Draconibacterium (Du et al., 2014); 3, Marinifilum (Na et al., 2009; Ruvira et al., 2013); 4, Prolixibacter (Holmes et al., 2007) +, Positive; 2, negative. All genera are non-motile and hydrolyse gelatin.

Characteristic 1 2 3 4

Origin Saltwater lake Marine Marine Marine Colony colour Pink Light pink to tawny Ivory or brownish ivory/unpigmented White Catalase ++ 22 Indole production 2 +++ Growth at: 4 uC 22 2 + 15 uC 22 2 + pH 5.5 2 + 2 + Hydrolysis of starch + 2 ++ Respiratory quinones MK-7 MK-7 MK-7 MK-7, MK-7(H6) DNA G+C content (mol%) 40.0 42.0 36 44.9

3474 International Journal of Systematic and Evolutionary Microbiology 64 Tangfeifania diversioriginum gen. nov., sp. nov.

Zhang et al., 2013) merely discovered the existence of the do so. Members of the genus Draconibacterium are also able novel 16S rRNA gene sequences with probable distribution to grow at pH 5.5 whereas strain G22T cannot. In addition, in different environments. To our knowledge, no bacterial G22T shows catalase activity, whereas members of the genus organisms belonging to a monophyletic lineage with Marinifilum do not. T strain G22 have been isolated previously. The EzTaxon-e T T Strain G22 also displays a number of differences from analysis revealed that strain G22 was most closely related to D. orientale, with 90.04 % sequence similarity, and more members of the genera Sunxiuqinia, Prolixibacter and distantly related to the representative members within the Mangrovibacterium. Major distinctive properties of the families Prolixibacteraceae and Marinilabiliaceae and the genus Sunxiuqinia include its inability to grow at 15 uC incertae sedis genus Marinifilum, with sequence similarities and at pH 5.5 as well as its ability to hydrolyse starch and gelatin. The genus Prolixibacter contains MK7 and MK- of 84.58–88.05 %. Such low levels of 16S rRNA gene T sequence similarity between strain G22T and its nearest 7(H6) as predominant menaquinones, whereas G22 dis- neighbours indicate that the isolate represents a novel plays only MK-7. Although the genus Mangrovibacterium genus. As shown in the NJ tree (Fig. 1), strain G22T is a contains MK-7 as a component of respiratory quinone, it also has MK-7(H2), MK-7(H6) and MK-9, which is distinct member of the class Bacteroidia and forms a distinct line of T descent within the family Draconibacteriaceae. The ML and from strain G22 . In addition, the members of the family Marinilabiliaceae display gliding motility, whereas this is not MP tree topologies were similar to the NJ tree topology T (Figs S2 and S3). observed for strain G22 . In addition to phylogenetic distance, strain G22T can In view of the results of our polyphasic chemotaxonomic be distinguished from its closest relatives by a number of analysis, we propose that strain G22T be assigned to a novel chemotaxonomic and phenotypic features (Table 1). It is species of a new genus within the family Draconibacteriaceae, pertinent to note that strain G22T can be readily distin- for which the name Tangfeifania diversioriginum gen. nov., guished from members of the genera Draconibacterium and sp. nov. is proposed. A detailed description of the morpho- Marinifilum by the ability of members of the latter two logical, physiological, biochemical and chemotaxonomic genera to produce indole, whereas strain G22T is unable to characteristics is given in the genus and species descriptions.

99 Sunxiuqinia elliptica LMG 25367T (GQ200190) 0.05 100 Sunxiuqinia faeciviva JCM 15547T (AB362263) DSM 27148T (JX983191) 96 Mangrovibacterium diazotrophicum 84 Meniscus glaucopis ATCC 29398T (NR_104837) 77 Prolixibacter bellariivorans JCM 13498T (AY918928) G22T (JQ683777) 97 Tangfeifania diversioriginum 68 Draconibacterium orientale FH5T (JQ683778)

100 Draconibacterium orientale SS4 (KF041475) 100 Marinifilum fragile JCM 15579T (FJ394546) 88 Marinifilum flexuosum DSM 21950T (HE613737) 82 Saccharicrinis fermentans DSM 9555T (M58766) Carboxylicivirga mesophila MEBiC 07026T (JQ672625) 94 T 100 Geofilum rubicundum JCM 15548 (AB362265) Natronoflexus pectinivorans DSM 24179T (GQ922844) 100 thermohalophila DSM 12881T (AJ418048) 100 Marinilabilia salmonicolor DSM 6480T (NR_104682) 100 Paraprevotella clara DSM 19731T (AB331896) 99 Prevotella melaninogenica ATCC 25845T (AY323525) Bacteroides fragilis ATCC 25285T (CR626927) 93 Porphyromonas asaccharolytica ATCC 25260T (L16490) T 74 Paludibacter propionicigenes DSM 17365 (CP002345) 68 Dysgonomonas gadei JCM 16698T (Y18530) Agarivorans albus LMG 21761T (AB076561)

Fig. 1. NJ tree generated using 16S rRNA gene sequences, showing the relationships of strain G22T to its closest relatives. Numbers at nodes represent bootstrap percentages (above 50 %) from 1000 replicates. Bar, 0.05 substitutions per nucleotide position. http://ijs.sgmjournals.org 3475 Q.-Q. Liu and others

Description of Tangfeifania gen. nov. b-glucuronidase, a-glucosidase, b-glucosidase, N-acetyl-b- glucosaminidase and a-mannosidase are absent. Cells are Tangfeifania (Tang.fei.fan9i.a. N.L. fem. n. Tangfeifania resistant to cefalotin, oxacillin, gentamicin, ciprofloxacin, named in honour of Tang Feifan (1897–1958), a Chinese kanamycin, norfloxacin, neomycin, amikacin, polymyxin microbiologist who was the first scientist to successfully B and trimethoprim, but sensitive to spectinomycin, isolate and culture Chlamydia trachomatis). erythromycin, midecamycin, acetylspiramycin, clindamy- Cells are Gram-stain-negative, straight or slightly curved cin, lincomycin and carbenicillin. The predominant cellular rods, non-endospore-forming, facultatively anaerobic and fatty acids are iso-C15 : 0, anteiso-C15 : 0, iso-C17 : 0 3-OH non-motile. The major polar lipids are phosphatidyletha- and iso-C15 : 0 3-OH. In addition to the major polar nolamine and three unknown lipids. The predominant lipids phosphatidylethanolamine and three unknown lipids, cellular fatty acids are iso-C15 : 0, anteiso-C15 : 0 and iso- moderate to minor amounts of glycolipid, aminolipid, two C17 : 0 3-OH. MK-7 is the main respiratory quinone. The unknown phospholipids and another unknown lipid are type species is Tangfeifania diversioriginum. present in the polar lipid profile. The type strain, G22T (5CICC 10587T5DSM 27063T), was Description of Tangfeifania diversioriginum isolated from Gahai, a saltwater lake located in Qinghai sp. nov. Province, China. The DNA G+C content of the type strain is 40.0 mol%. Tangfeifania diversioriginum (di.ver.si.o.ri9gi.num. L. adj. diversus diverse, different; origo -inis origin; N.L. gen. pl. n. diversioriginum of different origins). Acknowledgements Displays the following properties in addition to those This work was supported by the National Natural Science Foundation described for the genus. Cells are approximately 0.25– of China (31370057, 31290231), National Science and Technology 0.3 mm wide and 2.3–4.3 mm long. Surface colonies on Major Project of China (2013ZX10004217), and the China Ocean MA are pink, circular, convex, entire, smooth, opaque and Mineral Resources R & D Association (COMRA) Special Foundation about 0.5 mm in diameter after 7 days of incubation at (DY125-15-T-05). 33 uC. Catalase- and oxidase- positive. Growth occurs at 25–40 uC, pH 6.5–8.5 and in the presence of 1–8 % (w/v) References NaCl, with optimum growth between 33 and 35 uC, at pH 7.0–7.5 and with 2–4 % (w/v) NaCl. No growth occurs Al Masalma, M., Raoult, D. & Roux, V. (2009). Phocaeicola abscessus in the absence of NaCl. Nitrate is not reduced. Oxidation– gen. nov., sp. nov., an anaerobic bacterium isolated from a human fermentation test is weakly positive. Starch and Tween 80 brain abscess sample. Int J Syst Evol Microbiol 59, 2232–2237. are hydrolysed, but agar, alginate and CM-cellulose are not. Alain, K., Holler, T., Musat, F., Elvert, M., Treude, T. & Kru¨ ger, M. Results of API 20E assays are positive for tryptophan (2006). Microbiological investigation of methane- and hydrocarbon- deaminase, gelatinase and Simmons’ citrate utilization, discharging mud volcanoes in the Carpathian Mountains, Romania. Environ Microbiol 8, 574–590. but negative for ONPG, arginine dihydrolase, lysine decarboxylase, ornithine decarboxylase, urease, H S pro- Bowman, J. P. (2000). Description of Cellulophaga algicola sp. nov., 2 isolated from the surfaces of Antarctic algae, and reclassification of duction, indole production and Voges–Proskauer reaction. Cytophaga uliginosa (ZoBell and Upham 1944) Reichenbach 1989 as D-Fructose 6- phosphate, L-alanine and glucuronamide Cellulophaga uliginosa comb. nov. Int J Syst Evol Microbiol 50, 1861– are oxidized in Biolog GEN III assays. Acid is produced 1868. from D-arabinose, L-arabinose, D-ribose, D-xylose, L-xylose, Chang, Y. H., Cheng, T. W., Lai, W. J., Tsai, W. Y., Sun, C. H., Lin, L. H. D-fructose, L-sorbose, aesculin, turanose, D-lyxose, D- & Wang, P. L. (2012). Microbial methane cycling in a terrestrial mud tagatose and potassium 5-ketogluconate, but not from volcano in eastern Taiwan. Environ Microbiol 14, 895–908. glycerol, erythritol, D-adonitol, methyl b-D-xylopyranoside, Dong, X.-Z. & Cai, M.-Y. (2001). Determination of biochemical L-rhamnose, dulcitol, inositol, D-mannitol, D-sorbitol, characteristics. In Manual for Systematic Identification of General methyl a-D-mannopyranoside, methyl a-D-glucopyrano- , pp. 370–398. Edited by X.-Z. Dong & M.-Y. Cai. Beijing: side, amygdalin, arbutin, salicin, sucrose, trehalose, inulin, Science Press (in Chinese). melezitose, starch, glycogen, xylitol, gentiobiose, D-fucose, Du, Z.-J., Wang, Y., Dunlap, C., Rooney, A. P. & Chen, G.-J. (2014). D-arabitol, L-arabitol, potassium gluconate or potassium Draconibacterium orientale gen. nov., sp. nov., isolated from two distinct marine environments, and proposal of Draconibacteriaceae 2-ketogluconate. Acid production from D-galactose, D- fam. nov. Int J Syst Evol Microbiol 64, 1690–1696. glucose, D-mannose, N-acetylglucosamine, cellobiose, mal- Harris, J. K., Caporaso, J. G., Walker, J. J., Spear, J. R., Gold, N. J., tose, lactose, melibiose, raffinose and L-fucose is weakly Robertson, C. E., Hugenholtz, P., Goodrich, J., McDonald, D. & other positive in API 50CH assays. In assays with the API authors (2013). Phylogenetic stratigraphy in the Guerrero Negro ZYM system, alkaline phosphatase, esterase (C4), esterase hypersaline microbial mat. ISME J 7, 50–60. lipase (C8), leucine arylamidase, acid phosphatase, naph- Hiraishi, A., Ueda, Y., Ishihara, J. & Mori, T. (1996). Comparative thol-AS-BI-phosphohydrolase, a-galactosidase and b-fuco- lipoquinone analysis of influent sewage and activated sludge by high- sidase are present, but lipase (C14), valine arylamidase, performance liquid chromatography and photodiode array detection. cystine arylamidase, trypsin, chymotrypsin, b-galactosidase, J Gen Appl Microbiol 42, 457–469.

3476 International Journal of Systematic and Evolutionary Microbiology 64 Tangfeifania diversioriginum gen. nov., sp. nov.

Holmes, D. E., Nevin, K. P., Woodard, T. L., Peacock, A. D. & Lovley, Bacteroidetes isolated from sediment in a sea cucumber farm. Int J Syst D. R. (2007). Prolixibacter bellariivorans gen. nov., sp. nov., a sugar- Evol Microbiol 61, 2885–2889. fermenting, psychrotolerant anaerobe of the phylum Bacteroidetes, Ruvira, M. A., Lucena, T., Pujalte, M. J., Arahal, D. R. & Macia´ n, M. C. isolated from a marine-sediment fuel cell. Int J Syst Evol Microbiol 57, (2013). Marinifilum flexuosum sp. nov., a new Bacteroidetes isolated 701–707. from coastal Mediterranean Sea water and emended description of the Huang, X.-F., Liu, Y.-J., Dong, J.-D., Qu, L.-Y., Zhang, Y.-Y., Wang, genus Marinifilum Na et al., 2009. Syst Appl Microbiol 36, 155–159. F.-Z., Tian, X.-P. & Zhang, S. (2014). Mangrovibacterium diazotrophi- Sasser, M. (1990). Identification of bacteria by gas chromatography of cum gen. nov., sp. nov., a nitrogen-fixing bacterium isolated from a cellular fatty acids, MIDI Technical Note 101. Newark, DE: MIDI Inc. mangrove sediment, and proposal of Prolixibacteraceae fam. nov. Int J Swofford, D. L. (2002). PAUP*: Phylogenetic analysis using parsimony Syst Evol Microbiol 64, 875–881. (and other methods), version 4. Sunderland, MA: Sinauer Associates. Jorgensen, J. H., Turnidge, J. D. & Washington, J. A. (1999). Takai, K., Abe, M., Miyazaki, M., Koide, O., Nunoura, T., Imachi, H., Antibacterial susceptibility tests: dilution and disk diffusion methods. Inagaki, F. & Kobayashi, T. (2013). Sunxiuqinia faeciviva sp. nov., a In Manual of Clinical Microbiology, 7th edn, pp. 1526–1543. Edited by facultatively anaerobic organoheterotroph of the Bacteroidetes isolated P. R. Murray, E. J. Baron, M. A. Pfaller, F. C. Tenover & R. H. Yolken. from deep subseafloor sediment. Int J Syst Evol Microbiol 63, 1602– Washington, DC: American Society for Microbiology. 1609. Kim, O.-S., Cho, Y.-J., Lee, K., Yoon, S.-H., Kim, M., Na, H., Park, S.-C., Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013). Jeon, Y. S., Lee, J.-H. & other authors (2012). Introducing EzTaxon- MEGA6: molecular evolutionary genetics analysis version 6.0. Mol Biol e: a prokaryotic 16S rRNA gene sequence database with phylotypes Evol 30, 2725–2729. that represent uncultured species. Int J Syst Evol Microbiol 62, 716– 721. Thompson, J. D., Gibson, T. J., Plewniak, F., Jeanmougin, F. & Higgins, D. G. (1997). The CLUSTAL_X windows interface: flexible Liu, R., Zhang, Y., Ding, R., Li, D., Gao, Y. & Yang, M. (2009). strategies for multiple sequence alignment aided by quality analysis Comparison of archaeal and bacterial community structures in tools. Nucleic Acids Res 25, 4876–4882. heavily oil-contaminated and pristine soils. J Biosci Bioeng 108, 400– 407. Yang, S.-H., Seo, H.-S., Woo, J.-H., Oh, H.-M., Jang, H., Lee, J.-H., Kim, S.-J. & Kwon, K. K. (2014). Carboxylicivirga gen. nov. in Mesbah, M., Premachandran, U. & Whitman, W. B. (1989). Precise + the family Marinilabiliaceae with two novel species, Carboxylicivirga measurement of the G C content of deoxyribonucleic acid by high- mesophila sp. nov. and Carboxylicivirga taeanensis sp. nov., and performance liquid chromatography. Int J Syst Bacteriol 39, 159–167. reclassification of Cytophaga fermentans as Saccharicrinis fermentans Na, H., Kim, S., Moon, E. Y. & Chun, J. (2009). Marinifilum fragile gen. nov., comb. nov. Int J Syst Evol Microbiol 64, 1351–1358. gen. nov., sp. nov., isolated from tidal flat sediment. Int J Syst Evol Zhang, R., Wu, Q., Piceno, Y. M., Desantis, T. Z., Saunders, F. M., Microbiol 59, 2241–2246. Andersen, G. L. & Liu, W. T. (2013). Diversity of bacterioplankton in Qu, L., Zhu, F., Hong, X., Gao, W., Chen, J. & Sun, X. (2011). contrasting Tibetan lakes revealed by high-density microarray and Sunxiuqinia elliptica gen. nov., sp. nov., a member of the phylum clone library analysis. FEMS Microbiol Ecol 86, 277–287.

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