International Journal of Systematic and Evolutionary Microbiology (2014), 64, 1991–1997

DOI 10.1099/ijs.0.062695-0

Aquimarina pacifica sp. nov., isolated from seawater Zenghu Zhang, Tong Yu, Tingting Xu and Xiao-Hua Zhang Correspondence

College of Marine Life Sciences, Ocean University of China, Qingdao, PR China

Xiao-Hua Zhang [email protected]

A Gram-stain-negative, rod-shaped, non-flagellated, strictly aerobic bacterium with gliding motility, designated strain SW150T, was isolated from surface seawater of the South Pacific Gyre (396 199 S 1396 489 W) during the Integrated Ocean Drilling Program Expedition 329. Optimal growth occurred in the presence of 2–4 % (w/v) NaCl, at pH 7–8 and at 28–30 6C. The dominant fatty acids were iso-C15 : 0, iso-C17 : 0 3-OH, iso-C15 : 1 G, C16 : 1v6c and/or C16 : 1v7c and 10-methyl C16 : 0 and/or iso-C17 : 1v9c. The polar lipids of strain SW150T comprised phosphatidylethanolamine, three unknown polar lipids and one unknown aminolipid. The major respiratory quinone was MK-6. The DNA G+C content of strain SW150T was 33.8 mol%. Phylogenetic analysis based on 16S rRNA gene sequences showed that the novel strain was related most closely to Aquimarina macrocephali JAMB N27T and Aquimarina muelleri KMM 6020T with 97.8 and 96.8 % sequence similarities, respectively. The estimated DNA–DNA hybridization values were 21.00±2.33 % between strain SW150T and A. macrocephali JAMB N27T and 20.60±2.32 % between strain SW150T and Aquimarina megaterium XH134T. On the basis of polyphasic analyses, strain SW150T represents a novel species of the genus Aquimarina, for which the name Aquimarina pacifica sp. nov. is proposed. The type strain is SW150T (5JCM 18214T5CGMCC 1.12180T).

The genus Aquimarina, a member of the family Flavobacteriaceae, was proposed by Nedashkovskaya et al. (2005) for accommodation of heterotrophic, Gram-negative, aerobic, dark-yellow or brownish-coloured, gliding flavobacteria producing flexirubin-type pigments (Nedashkovskaya et al., 2005). Fourteen species of the genus Aquimarina have been recognized at the time of writing, all from marine environments: Aquimarina latercula (Lewin, 1969; Nedashkovskaya et al., 2005), Aquimarina muelleri (Nedashkovskaya et al., 2005), Aquimarina addita (Yi & Chun, 2011), Aquimarina longa (Yu et al., 2013) and Aquimarina megaterium (Yu et al., 2014) from sea water, Aquimarina brevivitae (Yoon et al., 2006; Nedashkovskaya et al., 2006) and Aquimarina macrocephali (Miyazaki et al., 2010) from sediment, Aquimarina intermedia (Nedashkovskaya et al., 2006) from a sea urchin, Aquimarina spongiae (Yoon et al., 2011) and Aquimarina amphilecti (Kennedy et al., 2014) from marine sponges, Aquimarina agarilytica (Lin et al., 2012) from the surface of the marine red alga Porphyra haitanensis, Aquimarina salinaria (Chen et al., Abbreviation: DDH, DNA–DNA hybridization. The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene sequence of Aquimarina pacifica SW150T is KC476291. The accession numbers for the draft genome sequences of strains SW150T, A. macrocephali JAMB N27T and A. megaterium XH134T are JACC00000000, JACA00000000 and JACB00000000, respectively. Two supplementary figures are available with the online version of this paper.

062695 G 2014 IUMS

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2012) from a saltpan, and Aquimarina mytili (Park et al., 2012) and Aquimarina gracilis (Park et al., 2013) from mussel. A novel bacterial strain, designated SW150T, was isolated from surface seawater in the South Pacific Gyre at station U1369 (39u 199 S 139u 489 W) during the Integrated Ocean Drilling Program Expedition 329. The aim of the present study was to determine the exact taxonomic position of strain SW150T using a polyphasic taxonomic approach. Seawater samples were spread on marine agar 2216 (MA; Becton Dickinson) and incubated at 28 uC. Strain SW150T was purified by streaking three times on MA. Cultures were maintained on MA plates at 28 uC and stocks were preserved in sterile 0.85 % (w/v) saline supplemented with 15 % (v/v) glycerol at 280 uC. A. muelleri LMG 22569T obtained from the Belgian Coordinated Collections of Microorganisms (BCCM), A. macrocephali JCM 15542T, A. latercula JCM 8515T and A. addita JCM 17106T obtained from the Japan Collection of Microorganisms (JCM), and A. longa SW024T and A. megaterium XH134T obtained from our own laboratory were used as reference strains, which were cultured as for strain SW150T [MA/marine broth 2216 (MB; BD), 28 uC], unless otherwise specified. Gram staining and flagellum staining were investigated using standard methods (Beveridge et al., 2007). Cell morphology was determined by transmission electron microscopy (JEM1200EX; JEOL) after cells had been negatively stained with 1991

Z. Zhang and others

1 % (w/v) phosphotungstic acid. Growth under anaerobic conditions was determined on MA after incubation in an anaerobic jar which was filled with nitrogen and with a packet of AneroPack-Anaero (Mitsubishi Gas Chemical) at 28 uC for 1 month. Salt tolerance was investigated by using synthetic marine ZoBell broth (5 g Bacto peptone, 1 g yeast extract and 0.1 g FePO4 in 1 l modified artificial seawater; Lyman & Fleming, 1940) supplied with various concentrations of NaCl (0–15 %, w/v, at intervals of 1 %). In the modified artificial seawater, all of the Na+ was replaced by appropriate K+. Growth with NaCl as the sole salt was investigated on MA (distilled water instead of seawater) supplemented with 0–15 % (w/v) NaCl. The temperature range for growth was determined on MA plates incubated at 8–42 uC (8, 16, 28, 30, 34, 37 and 42 uC) for 1 week and at 0 and 4 uC on MA for 4 weeks. The pH range for growth was determined on MA at pH 2–10 at intervals of 1 pH unit. The presence of gliding motility and the production of flexirubin-type pigments were investigated using the methods described by Bernardet et al. (2002). In brief, bacterial strains were cultured on MA at 28 uC for 3 days, and the absorption spectrum of pigments extracted with acetone/ methanol (7 : 2, v/v) was determined at 300–700 nm with a UV–visible spectrophotometer (TU-1810; Beijing Purkinje General Instrument) (Tindall et al., 2007). Standard protocols (Tindall et al., 2007) were used to assess catalase and oxidase activities, degradation of casein, urea, starch, gelatin and Tweens 20, 40 and 80, agar corrosion, nitrate reduction and H2S production from thiosulfate with the modification that sterile seawater was used. DNase activity was examined by using DNase agar (Qingdao Hope Bio-technology) with sterile seawater, according to the manufacturer’s instructions. Degradation of chitin was examined on chitin agar with sterile seawater (Hsu & Lockwood, 1975). Utilization of substrates as sole carbon and energy sources was tested for 2 weeks in artificial seawater supplemented with 0.2 % (w/v) NaNO3 and 1 % (w/v) of the substrate. Activities of constitutive enzymes and other physiological properties were determined after growth on MA at 28 uC for 2 days by using API 20E, API 20NE, API 50CH and API ZYM strips (bioMe´rieux) according to the manufacturer’s instructions except that sterile seawater was used to prepare the inocula. Susceptibility to antibiotics was investigated on MA plates by using discs (Hangzhou Microbiology Reagent) containing different antibiotics. The detailed morphological, physiological and biochemical characteristics of strain SW150T are given in the species description, Table 1 and Fig. 1.

RTSBA6 6.10 database (Sasser, 1990). The respiratory quinones of strain SW150T were extracted with chloroform/ methanol (2 : 1, v/v), separated by TLC and identified by HPLC as described by Collins (1994). Polar lipids were analysed by using standard procedures (Minnikin et al., 1984). Extracted lipids were separated by two-dimensional TLC and identified by spraying with appropriate detection reagents. Spraying with 5 % ethanolic molybdophosphoric acid followed by charring at 150 uC revealed the presence of all lipids. Ninhydrin spray was applied to determine most of the lipids with free amino groups while spraying the plate with the lipid phosphate reagent of Dittmer and Lester after the ninhydrin spray revealed the presence of phospholipids. Spraying with a-naphthol-sulphuric acid determined the presence of glycolipids. The periodate–Schiff reagent spray was applied to reveal lipids with a terminal CH2OH group, such as glycerol (Minnikin et al., 1984). The G+C content of the chromosomal DNA of strain SW150T was determined according to the methods described by Mesbah & Whitman (1989) using reversed-phase HPLC. The dominant cellular fatty acids of strain SW150T were iso-C15 : 0 (19.6 %), isoC17 : 0 3-OH (13.8 %), iso-C15 : 1 G (10.7 %), C16 : 1v6c and/or C16 : 1v7c (8.3 %) and 10-methyl C16 : 0 and/or iso-C17 : 1v9c (7.7 %). The fatty acid profile of strain SW150T was essentially similar to those of the six reference strains except for the proportions of some fatty acids (Table 2). In accordance with other members of the family Flavobacteriaceae, the major respiratory quinone of strain SW150T was MK-6. The polar lipids were phosphatidylethanolamine, three unknown polar lipids and one unknown aminolipid (Fig. S1, available in the online Supplementary Material), which was very similar to those of A. macrocephali JCM 15542T, A. muelleri LMG 22569T, A. megaterium XH134T and A. longa SW024T, with only minor differences in the proportions of some lipids. The DNA G+C content of strain SW150T was 33.8 mol%, a value in the range of those for recognized species of the genus Aquimarina (Table 1).

For fatty acid analysis, cell masses of strains SW150T, A. longa SW024T, A. megaterium XH134T, A. muelleri LMG 22569T, A. macrocephali JCM 15542T, A. latercula JCM 8515T and A. addita JCM 17106T were obtained after cultivation on MA at 28 uC for 2–3 days when all of the communities reached the late exponential stage of growth. Extraction of fatty acid methyl esters and separation by GC were performed by using the Instant FAME method of the Microbial Identification System (MIDI) version 6.1 and the

For 16S rRNA gene sequencing, DNA was extracted from strain SW150T and purified using standard methods (Ausubel et al., 1995). The 16S rRNA gene was amplified by PCR using two universal primers (B8F, 59-AGAGTTTGATCCTGGCTCAG-39; B1510, 59-GGTTACCTTGTTACGACTT-39). The PCR product was purified using the TIANgel Midi Purification kit (TIANGEN Biotech), cloned into pUCmT vector (TaKaRa) and sequenced using an automated DNA sequencer (model ABI3730; Applied BioSystems) at BGI, Qingdao, China. The nearly complete 16S rRNA gene sequence (1483 nt) of strain SW150T was submitted to GenBank, and the identification of phylogenetic neighbours and the calculation of pairwise 16S rRNA gene sequence similarities were achieved using the EzTaxon-e server (http:// eztaxon-e.ezbiocloud.net/; Kim et al., 2012). Sequences were aligned using CLUSTAL X1.8 (Thompson et al., 1997). Phylogenetic trees were reconstructed using the neighbourjoining and maximum-likelihood methods with Kimura two-state parameter model analyses (Kimura, 1980) implemented in the program MEGA version 5 (Tamura et al., 2011).

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International Journal of Systematic and Evolutionary Microbiology 64

Aquimarina pacifica sp. nov.

Table 1. Differential characteristics of strain SW150T and other members of the genus Aquimarina Strains: 1, Aquimarina pacifica sp. nov. SW150T; 2, A. macrocephali JCM 15542T; 3, A. muelleri LMG 22569T; 4, A. megaterium XH134T; 5, A. latercula JCM 8515T; 6, A. longa SW024T; 7, A. addita JCM 17106T. All data are from this study and Yu et al. (2013, 2014) except for DNA G+C content and cell size, which are from the original descriptions. +, Positive reaction; ”, negative reaction; W, weakly positive reaction. Characteristic Colony colour Cell characteristics Width (mm) Length (mm) Gliding motility Range for growth Temperature (uC) NaCl (%) Flexirubin-type pigments Nitrate reduction Hydrolysis of: Aesculin Agar Casein Chitin Starch Tween 20 Tween 40 Tween 80 Utilization of: D-Mannose Leucine Lysine Citrate API ZYM results Cystine arylamidase Trypsin a-Chymotrypsin Naphthol-AS-BIphosphohydrolase a-Glucosidase Susceptibility to: Ampicillin Carbenicillin Cefuroxime Cefoperazone Erythromycin Lincomycin Minocycline Norfloxacin Piperacillin Rocephin Tetracycline Vancomycin Maximum absorption of pigments (nm) DNA G+C content (mol%)

1

2

3

4

5

6

7

Orange

Orange

Yellow–brown

Orange

Orange–red

Yellow

Orange

0.2 2.5–4.5 +

0.4–0.6 3.0–25.5 +

0.3–0.5 5–7 +

0.5 5.4–77.8 +

0.3–0.4 1–5 –

0.3 3.0–66.0 +

0.3–0.5 1.4–2.2 –

8–30 2–4 – –

8–30 1–4 + +a*

4–34 1–7 + +b

8–37 2–4 + +

4–34 1–5 + +b

4–30 3–5 – +

4–30 3–7 – –c

+ – – – –

– – + + + + + +

– – – + + + +

+ + –b – +b + +

W

– – –a + + – – –

W

W

– – – + – + + +

+ – – + + + + +

– – – –

– – – –

– + + –

– + + +

– – – –

+ – – –

– – – –



– –



W

W

W

W

W

W

W

W

W

W

W

W

+

– – – +

– –c +c +

W





W





– – – – + + – +

– – – – + – – +

– + 453; 479

+ + – + 506

30.8

35

W



W

– + +

b

+ – – – + + – + + + – + 477

W

+ + + – + + – + +

– – + 483a

W

W

+ – 472b

– + 473

+ – – + + + + + + + – + 503d

33.8

33.1

31–33

32.4

34

W

W

– – – W W

– + + W

+ + + – + + – W

*Data different from: a, Miyazaki et al. (2010); b, Nedashkovskaya et al. (2005); c, Yi & Chun (2011); d, Nedashkovskaya et al. (2006).

http://ijs.sgmjournals.org

1993

Z. Zhang and others

(a)

calculator (GGDC2.0) (Auch et al., 2010a, b; Meier-Kolthoff et al., 2013).

(b)

Phylogenetic analysis of the 16S rRNA gene sequence indicated that strain SW150T had the highest pairwise similarity with A. macrocephali JAMB N27T (97.8 %), A. muelleri KMM 6020T (96.8 %) and A. megaterium XH134T (96.6 %). The similarities between strain SW150T and other species of the genus Aquimarina were in the range 94.6– 97.8 %. Phylogenetic analysis based on neighbour-joining (Fig. 2), maximum-likelihood (Fig. S2) and maximumparsimony algorithms showed that strain SW150T formed a distinct cluster with the genus Aquimarina. The estimated DDH values calculated using GGDC2.0 with the alignment method BLAST+ were 21.00±2.33 % between strains SW150T and A. macrocephali JAMB N27T and 20.60±2.32 % between strains SW150T and A. megaterium XH134T.

Fig. 1. Transmission electron micrographs of negatively stained cells of strain SW150T cultured on MA at 28 6C for 48 h. Bars, 500 nm (a), 200 nm (b).

The major features of strain SW150T including chemotaxonomic characteristics and DNA G+C content were in line with those of the six reference strains of the genus Aquimarina and the phylogenetic analysis based on 16S rRNA gene sequences showed that strain SW150T belongs to the genus Aquimarina. However, the low levels of sequence

In each case, bootstrap values were calculated based on 1000 replicates. Estimated DNA–DNA hybridization (DDH) values were analysed using the genome-to-genome distance

Table 2. Cellular fatty acid compositions of strain SW150T and other members of the genus Aquimarina Strains: 1, Aquimarina pacifica sp. nov. SW150T; 2, A. macrocephali JCM 15542T; 3, A. muelleri LMG 22569T; 4, A. megaterium XH134T; 5, A. latercula JCM 8515T; 6, A. longa SW024T; 7, A. addita JCM 17106T. All data are from this study and Yu et al. (2013, 2014). TR, Traces (,1 %); –, not detected. Values are percentages of the total fatty acids. Fatty acids amounting to ,1 % of the total in all strains are not shown. Fatty acid iso-C13 : 0 C13 : 1 at 12-13 C14 : 0 C15 : 0 2-OH iso-C15 : 0 iso-C15 : 1 G C15 : 1v6c iso-C15 : 0 3-OH C16 : 0 iso-C16 : 0 anteiso-C16 : 0 iso-C16 : 1 H iso-C16 : 1 G C16 : 0 3-OH iso-C16 : 0 3-OH C17 : 1v6c C17 : 1v8c iso-C17 : 0 3-OH C20 : 1v9c Summed features* C16 : 1v6c and/or C16 : 1v7c 10-Methyl C16 : 0 and/or iso-C17 : 1v9c

1

2

3

5.5

1.6

1.1

TR

3.8

TR

TR

TR

TR

TR

TR

TR

5.6 3.4 1.5

TR

1.0

TR

TR

TR

TR

TR

TR

TR

TR

19.6 10.7 4.0 5.4 3.7 4.5 1.5 – 2.1

20.4 6.6

26.7 8.3

1.3 22.5 11.8

1.5 24.2 9.1 5.2 4.8 1.8 2.1 – 1.6 – 1.1 1.1 2.0 2.0 13.9

TR

4

5

TR

TR

10.0 1.1 – –

TR

5.7 4.8 7.6 – 3.2 – 1.7

– –

33.5 12.6 – 7.3 5.4 3.9 – – 1.1 1.4

TR

TR

TR

TR

1.2 2.7 13.8

TR

TR

TR

TR

TR

TR

TR

TR

8.3 7.7

TR

TR

5.4 2.4 8.5 2.6 – 3.1 2.2 1.3 2.9

6

34.8 10.5 TR

7.5 1.4 3.3 – 1.7 – TR TR TR TR

7.7 1.7

18.9 –

8.6 –

12.0 –

15.9 –

15.1 7.5

11.6 13.9

11.7 7.0

9.0 6.3

4.3 9.8

7

TR

10.6 12.3

*Summed features are groups of two or three fatty acids that are treated together for the purpose of evaluation in the MIDI system and include both peaks with discrete equivalent chain lengths (ECLs) as well as those where the ECLs are not reported separately, as indicated by MonteroCalasanz et al. (2013). 1994

International Journal of Systematic and Evolutionary Microbiology 64

Aquimarina pacifica sp. nov.

90

Aquimarina longa SW024T (JN118549) Aquimarina pacifica SW150T (KC476291)

96

Aquimarina macrocephali JAMB N27T (AB517144) Aquimarina megaterium XH134T (KC476292)

0.01

Aquimarina addita JC2680T (HM475137) ‘Aquimarina litoralis’ CNURIC011 (FJ490365) Aquimarina latercula ATCC 23177T (M58769)

86 95

70

Aquimarina amphilecti 92VT (JX050189) Aquimarina intermedia KMM 6258T (AM113977) Aquimarina mytili PSC33T (HM998910) Aquimarina salinaria antisso-27T (FJ882164)

91

Aquimarina gracilis PSC32T (HM998909) Aquimarina spongiae A6T (FJ348470) Aquimarina agarilytica ZC1T (FJ750453) Dokdonia eikasta PMA-26T (AB198088) Dokdonia donghaensis DSW-1T (DQ003276) 100 Dokdonia genika Cos-13T (AB198086) 98

99

99

100

Lacinutrix algicola AKS293T (DQ167238) Lacinutrix mariniflava AKS432T (DQ167239)

Algibacter mikhailovii LMG 23988T (AM491809) 100

91

100

Lacinutrix copepodicola DJ3T (AY694001) Algibacter lectus KMM 3902T (Y187689)

99

100

95

Aquimarina muelleri KMM 6020T (AY608406)

Formosa agariphila KMM 3901T (AY187688) Formosa algae KMM 3553T (AY228461)

Bizionia argentinensis JUB59T (EU021217) Bizionia algoritergicola APA-1T (AY694003) 100 91 Bizionia echini KMM 6177T (FJ716799) Kordia algicida OT-1T (AY195836) Kordia aquimaris CC-AMZ-301T (JX235671) Cellulophaga fucicola NN015860T (AJ005973) Cellulophaga pacifica KMM 3664T (AB100840) Cytophaga fermentans ATCC 19072T (M58766)

Fig. 2. Neighbour-joining phylogenetic tree based on 16S rRNA gene sequences showing the phylogenetic positions of strain SW150T, the type strains of other species of the genus Aquimarina and representatives of some other related members of the family Flavobacteriaceae. Percentage bootstrap values above 70 (1000 replicates) are shown at branch nodes. Cytophaga fermentans ATCC 19072T (M58766) was used as an outgroup. Bar, 0.01 substitutions per nucleotide position.

similarity to the type strains of recognized species of the genus Aquimarina and the estimated DDH values with strains A. macrocephali JAMB N27T and A. megaterium XH134T below the 70 % cut off point for recognition of genomic species (Wayne et al., 1987) implied that strain SW150T may represent a novel species. Moreover, a number of phenotype characteristics (Table 1), namely inability to reduce nitrate, absence of flexirubin-type pigments and inability to grow well in MB, are the most obvious differences between strain SW150T and the recognized species of genus Aquimarina. On the basis of phenotypic characteristics and phylogenetic inference, strain SW150T is assigned to the genus Aquimarina as a representative of a novel species, for which the name Aquimarina pacifica sp. nov. is proposed. http://ijs.sgmjournals.org

Description of Aquimarina pacifica sp. nov. Aquimarina pacifica (pa.ci9fi.ca. L. fem. adj. pacifica peaceful, referring to the Pacific Ocean from where the type strain was isolated). Cells are Gram-stain-negative, strictly aerobic rods, approximately 2.5–4.5 mm in length and 0.2 mm in width, nonflagellated and motile by gliding. Colonies on MA are orange, slightly transparent, convex, circular with non-entire margins and 0.5–1.0 mm in diameter after 2–3 days at 28 uC. Does not grow well in MB. Growth occurs at 8–30 uC. NaCl alone does not support growth; requires seawater or artificial seawater for growth. Growth occurs in MB with 2–4 % (w/v) NaCl. The pH range for growth is pH 7–8. Pigments are formed with maximum absorption at 477 nm, but flexirubin-type pigments are not formed. Oxidase and 1995

Z. Zhang and others

catalase are positive. Acetoin, H2S and indole are not produced. Nitrate is not reduced. Gelatin, DNA, aesculin and Tweens 20 and 80 are hydrolysed (Tweens 20 and 80 are weakly hydrolysed), but Tween 40, starch, aga, casein, chitin and urea are not hydrolysed. Arginine dihydrolase, galactosidase, lysine decarboxylase, ornithine decarboxylase and tryptophan deaminase are absent. Acid is not produced in the API 50CH system. According to carbon source experiments, mannose, sucrose, a-lactose, fructose, leucine, xylose, glucose, ornithine, lysine, raffinose, rhamnose and citrate cannot be utilized. In the API ZYM system, alkaline phosphatase, esterase (C4), esterase lipase (C8), leucine arylamidase, valine arylamidase, acid phosphatase, naphtholAS-BI-phosphohydrolase and a-glucosidase activities are present, trypsin activity is weakly present, but lipase (C14), cystine arylamidase, a-fucosidase, chymotrypsin, N-acetyl-bglucosaminidase, b-glucosidase, b-glucuronidase, a-galactosidase, glucuronidase and a-mannosidase activities are absent. Resistant to benzylpenicillin G, cephalosporin V, cefuroxime, polymyxin B, gentamicin, kanamycin, neomycin, tetracycline, minocycline, cefoperazone and streptomycin. Susceptible to ampicillin, piperacillin, erythromycin, vancomycin, norfloxacin, Rocephin, furazolidone, clindamycin, chloramphenicol, rifampicin and lincomycin, and intermediately susceptible to ciprofloxacin. The major respiratory quinone is MK-6. The dominant fatty acids are iso-C15 : 0, iso-C17 : 0 3-OH, iso-C15 : 1 G, C16 : 1v6c and/or C16 : 1v7c and 10-methyl C16 : 0 and/or iso-C17 : 1v9c. The polar lipid profile comprises phosphatidylethanolamine, three unknown polar lipids and one unknown aminolipid. The type strain, SW150T (5JCM 18214T5CGMCC 1.12180T), was isolated from surface seawater of the South Pacific Gyre. The DNA G+C content of the type strain is 33.8 mol%.

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Acknowledgements

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This research used samples provided by the Integrated Ocean Drilling Program (IODP). We thank all of the crew and technical staff on the JOIDES Resolution during the IODP Expedition 329 for their great efforts and help in sample collection. This work was supported by the National Natural Science Foundation of China (no. 41276141), the National High Technology Research & Development Program of China (863 Programs, no. 2012AA091605) and the China Ocean Mineral Resources Research & Development Association (COMRA, no. DY125-15-R-03).

Meier-Kolthoff, J. P., Auch, A. F., Klenk, H. P. & Go¨ker, M. (2013).

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A Gram-stain-negative, rod-shaped, non-flagellated, strictly aerobic bacterium with gliding motility, designated strain SW150(T), was isolated from su...
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