Report of 39 Unrecorded Bacterial Species in Korea Belonging to Gammaproteobacteria

Journal24 of Species Research 7(1):24-35, 2018Journal of Species Research Vol. 7, No. 1

Report of 39 unrecorded bacterial species in Korea belonging to Gammaproteobacteria

Min-Kyeong Kim1, Jisun Park1, Bo-Ram Yun1, Jin-Woo Bae2, Chang-Jun Cha3, Jang-Cheon Cho4, Wan-Taek Im5, Kwang Yeop Jahng6, Che Ok Jeon7, Kiseong Joh8, Wonyong Kim9, Soon Dong Lee10, Chi Nam Seong11, Hana Yi12 and Seung-Bum Kim1,*

1Department of Microbiology and Molecular Biology, Chungnam National University, Daejeon 34134, Republic of Korea 2Department of Biology, Kyung Hee University, Seoul 02447, Republic of Korea 3Department of Biotechnology, Chung-Ang University, Anseong 17546, Republic of Korea 4Department of Biological Sciences, Inha University, Incheon 22212, Republic of Korea 5Department of Biotechnology, Hankyoung National University, Anseong 17579, Republic of Korea 6Department of Biological Sciences, Chonbuk National Universty, Jeonju 54896, Republic of Korea 7Department of Life Science, Chung-Ang University, Seoul 06974, Republic of Korea 8Department of Biotechnology, Hankuk University of Foreign Studies, Yongin 17035, Republic of Korea 9Department of Microbiology, Chung-Ang University College of Medicine, Seoul 06974, Republic of Korea 10Department of Science Education, Jeju National University, Jeju 63243, Republic of Korea 11Department of Biology, Sunchon National University, Suncheon 57922, Republic of Korea 12School of Biosystem and Biomedical Science, Korea University, Seoul 02841, Republic of Korea

*Correspondent: [email protected]

During a series of extensive surveys of prokaryotic species diversity in Korea, bacterial strains belonging to Gammaproteobacteria were isolated from various sources of aquatic and terrestrial environments. A total of 39 isolates were obtained, which represented 39 unrecorded species in Korea belonging to 20 genera in 12 families. Enterobacteriaceae was the largest family, as eight species were assigned, which was followed by Moraxellaceae (6 species) and Pseudomonadaceae (5 species). At the genus level, Marinobacter (6 species), and Pseudomonas (5 species) were the main genera, and at least two species were obtained for Acinetobacter (3 species), Psychrobacter (3 species), Shewanella (2 species), Dickeya (2 species), Salinivibrio (2 species), Vibrio (2 species) and Rhodanobacter (2 species). The detailed description of each unrecorded species is provided. Keywords: 16S rRNA, Enterobacteriaceae, Gammaproteobacteria, Marinobacter, unrecorded species

Introduction are also diverse, including terrestrial and aquatic environments, environments with extreme physicochemical The phylum Proteobacteria is currently the largest conditions, and animal and plant hosts. known bacterial group, and among the classes of Pro- The diverse environments of Korea have contributed to teobacteria, Gammaproteobacteria is comprised of the the biodiversity of Gammaproteobacteria, as a number of broadest groups of proteobacteria, including 21 orders, unrecorded gammaproteobacterial species have already 60 families and 394 genera as of August 2017 (Euzéby, been reported in previous studies (Kim et al., 2011; Choi 2017). et al., 2015; Jung et al., 2016; Lim et al., 2016). Mem- Gammaproteobacteria encompasses anoxygenic pho- bers of the families Moraxellaceae, Pseudomonadaceae, totrophs, various chemolithotrophs and fermenters as and Enterobacteriaceae were consistently recovered in well as aerobic or anaerobic respiratory heterotrophs, all studies, and Xanthomonadaceae, Alteromonadaceae, thus exhibiting virtually all types of metabolism found Pseudoalteromonadaceae, Vibrionaceae were also fre- among prokaryotes (Brenner et al., 2005). Their habitats quently recovered constituents in those studies. February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 25

In the following series of extensive surveys on the strains represented 39 unrecorded species in Korea be- prokaryotic diversity in Korea, novel strains belonging longing to 20 genera of 12 families. Enterobacteriaceae to the class Gammaproteobacteria of the phylum Pro- was the largest family, as eight species were assigned, teobacteria were isolated from various environmental which was followed by Moraxellaceae (6 species), sources, and their taxonomic properties were analyzed. Pseudomonadaceae (5 species) and Vibrionaceae (4 spe- As a result, a total of 39 unrecorded species of Gam- cies). One or two species were recovered for the remain- maproteobacteria were identified, and their taxonomic ing nine families (Table 1). At the genus level, Marino- properties are presented. bacter (6 species), and Pseudomonas (5 species) were the main genera, and at least two species were obtained for Acinetobacter (3 species), Psychrobacter (3 species),

Materials and Methods Shewanella (2 species), Dickeya (2 species), Salinivibrio (2 species), Vibrio (2 species) and Rhodanobacter (2 spe- A total of 39 bacterial strains assigned to the class cies). Single species were recovered for the remaining Gammaproteobacteria were isolated from diverse envi- 11 genera (Table 1). The electron microscopic images of ronmental samples collected from sediment, tidal flats, the isolates are provided in Fig. 1, and the phylogenetic soil, seawater, freshwater, brown alga, mealworm, solar relationship between the isolates and closely related spe- saltern, plants and sewage treatment plants (Table 1). cies are presented in Figs. 2-5. The detailed description The samples were processed or treated separately, dilut- of each unrecorded species is provided below. ed and spread onto diverse culture media R2A, marine agar (MA), tryptic soy agar (TSA) or nutrient agar (NA) Description of Acinetobacter gandensis 4003 (BD), and incubated at 25-30°C for 2-5 days. The desig- nated strain IDs, sources of isolation, culture media, and Cells are Gram-stain-negative, non-flagellated, non- incubation conditions are provided in the description pigmented and coccoid-shaped. Colonies are irregular, and Table 1. All strains were purified as single colonies entire, smooth, raised and white colored after incubation and stored as 10-20% glycerol suspension at -80°C as for 3 days on R2A at 30℃. Positive for nitrate reduc- well as lyophilized ampoules. tion, indole production, glucose fermentation, urease, Colony characteristics of the isolates were observed esculin hydrolysis and β-galactosidase, but negative on the same agar media for cultivation. Cellular mor- for arginine dihydrolase and gelatin hydrolysis in API phology and cell size were examined using either a 20NE. D-glucose, L-arabinose, D-mannose, D-manni- transmission electron microscope or a scanning electron tol, N-acetyl-glucosamine, D-maltose, potassium gluco- microscope. Gram staining was performed using the nate, capric acid, malic acid, trisodium citrate and phe- standard procedures. Biochemical characteristics were nylacetic acid are utilized as sole carbon sources, but not = tested by using API 20NE galleries (bioMérieux) ac- adipic acid. Strain 4003 ( NIBRBAC000498564) was cording to the manufacturer’s instructions. isolated from a sediment sample, Seoul, Korea. The extraction of genomic DNA, PCR amplification of 16S rRNA gene and sequencing were performed Description of Raoultella electrica 4231 using the procedures as described previously (Choi et Cells are Gram-stain-negative, non-flagellated, non- al., 2015; Jung et al., 2016; Um et al., 2016). The 16S pigmented and rod-shaped. Colonies are circular, en- rRNA gene sequences of the strains were compared tire, smooth, flat and white colored after incubation for with the reference strains using the EzBioCloud (Kim et 2 days on R2A at 30℃. Positive for nitrate reduction, al., 2012), and their 16S rRNA sequences were aligned glucose fermentation, urease, esculin hydrolysis and β- using EzEditor2 (Jeon et al., 2014). Phylogenetic trees galactosidase, but negative for indole production, argi- were generated by Jukes-Cantor distance model (Jukes nine dihydrolase and gelatin hydrolysis in API 20NE. and Cantor, 1969) and neighbor-joining method (Saitou D-glucose, L-arabinose, D-mannose, D-mannitol, N- and Nei, 1987) using MEGA 7.0 (Kumar et al., 2016). acetyl-glucosamine, D-maltose, potassium gluconate, The phylogenetic trees were evaluated using the boot- adipic acid, malic acid and trisodium citrate are utilized strap analysis (Felsenstein, 1985) based on 1,000 resam- as sole carbon sources, but not capric acid and pheny- pled dataset. lacetic acid. Strain 4231 (=NIBRBAC000498567) was isolated from a sediment sample, Seoul, Korea.

Results and Discussion Description of Dickeya zeae 7026 A total of 39 bacterial strains belonging to the class Cells are Gram-stain-negative, flagellated, non-pig- Gammaproteobacteria were isolated (Table 1). The mented and rod-shaped. Colonies are circular, entire, 26 Journal of Species Research Vol. 7, No. 1

conditions I ncubation 30℃, 2 days 20℃, 3 days 25℃, 2 days 30℃, 3 days 30℃, 2 days 25℃, 4 days 25℃, 3 days 37℃, 2 days 37℃, 2 days 37℃, 2 days 25℃, 3 days 20℃, 3 days 25℃, 1 day 25℃, 1 day 25℃, 1 day 30℃, 2 days 20℃, 3 days 25℃, 3 days 25℃, 2 days 25℃, 3 days 30℃, 2 days 20℃, 3 days 25℃, 1 day 25℃, 5 days 30℃, 2 days 30℃, 2 days 25℃, 3 days 30℃, 4 days 30℃, 2 days 30℃, 2 days 25℃, 1 day 30℃, 2 days 25℃, 1 day 30℃, 2 days 25℃, 4 days 20℃, 3 days 25℃, 3 days 25℃, 3 days 25℃, 2 days (p H 9)

with A SW with A SW (p H 8)

Medium M A TS A TS A TS A R 2 A TS A M A M A M A M A M A M A M A M A M A M A R 2 A R 2 A M A M A M A M A M A R 2 A R 2 A R 2 A TS A M A R 2 A L B R 2 A M A R 2 A R 2 A M A R 2 A R 2 A TS A M A I solation source P lant root Sea water Sea water C ave F reshwater Sea water Saltern Saltern Saltern fulvellum Sargassum Tidal flat Tidal unicinctus Urechis flat Tidal flat Tidal Soil flat Tidal Ginseng field Sewage treatment plant Ecklonia cava Sewage treatment plant Tidal flat Tidal flat Tidal Zostera marina Sediment Sediment Tree Tidal flat Tidal Sewage treatment plant Sediment Tenebrio molitor Tenebrio Sediment Tidal flat Tidal Sediment F reshwater P lant root Sea water Wastewater Sea water Sea water

100 100 100 100 100 100 100 (%) 98.8 98.8 99.7 99.0 99.6 99.3 99.5 99.3 99.9 99.9 99.4 98.9 99.1 99.7 99.1 99.7 99.2 99.9 99.2 99.9 99.9 99.4 99.7 99.2 99.2 98.7 98.5 99.9 99.6 99.8 99.9 98.7 Similarity . vallismortis indologenes subsp subsp. Most closely related species Pseudomonas alcaliphila Pseudomonas guineae Pseudomonas salina Pseudomonas cuatrocienegasensis Aeromonas allosaccharophila Aeromonas Haemophilus piscium Salinivibrio costicola Salinivibrio sharmensis Vibrio ruber Vibrio fortis Vibrio Marinobacter hydrocarbonoclasticus Marinobacter oulmenensis Marinobacter sediminum Marinobacter guineae taiwanensis Microbulbifer mangrovi Microbulbifer Rhodanobacter spathiphylli Rhodanobacter glycinis Pseudoalteromonas lipolytica Pseudoalteromonas Xanthomonas sacchari Shewanella algae Shewanella algidipiscicola Arenicella chitinivorans Arenicella Dickeya zeae Dickeya chrysanthemi Erwinia billingiae Escherichia fergusonii Pantoea stewartii Raoultella electrica Serratia quinivorans Yokenella regensburgei Yokenella Marinobacterium nitratireducens Acinetobacter gandensis Acinetobacter junii Acinetobacter radioresistens Psychrobacter marincola Psychrobacter faecalis Psychrobacter piscatorii Psychrobacter Pseudomonas punonensis NI B R I D NI B R AC 000498551 NI B R AC 000498618 NI B R AC 000498619 NI B R AC 000498655 NI B R AC 000498636 NI B R AC 000498616 NI B R AC 000498456 NI B R AC 000498458 NI B R AC 000498457 NI B R AC 000498472 NI B R AC 000498542 NI B R AC 000498520 NI B R AC 000498527 NI B R AC 000498530 NI B R AC 000498497 NI B R AC 000498541 NI B R AC 000498590 NI B R AC 000498406 NI B R AC 000498484 NI B R AC 000498425 NI B R AC 000498534 NI B R AC 000498525 NI B R AC 000498478 NI B R AC 000498574 NI B R AC 000498576 NI B R AC 000498453 NI B R AC 000498660 NI B R AC 000498426 NI B R AC 000498567 NI B R AC 000498486 NI B R AC 000498575 NI B R AC 000498538 NI B R AC 000498564 NI B R AC 000498630 NI B R AC 000498546 NI B R AC 000498632 NI B R AC 000498473 NI B R AC 000498615 NI B R AC 000498504 Strain I D I M CC 25652 UL 498-1 UL 506 Y C 7-24 KB012 UL 222 H M F 8002 H M F 8044 H M F 8004 S F D22 I M CC 25643 LP B0132 LP B0156 LP B0159 CAU 1046 I M CC 25642 Gsoil 3009 Y H Y1 EC 2D12 S N 5 I M CC 25635 LP B0153 Z O D21 7026 9023 H M F 7415 B E 6-1 S N 10 4231 GM22 7034 I M CC 25639 4003 DS041 I M CC 25647 L GGG06 PO B9 UL 158 CAU 1335 Genus Aeromonas Haemophilus Salinivibrio Vibrio Marinobacter Rhodanobacter Pseudoalteromonas Xanthomonas Shewanella Arenicella Dickeya Erwinia Escherichia Pantoea Raoultella Serratia Yokenella Marinobacterium Acinetobacter Psychrobacter Pseudomonas F amily Aeromonadaceae Vibrionaceae Alteromonadaceae Rhodanobacteraceae Pseudoalteromonadaceae Xanthomonadaceae Shewanellaceae Arenicellaceae Enterobacteriaceae Oceanospirillaceae Moraxellaceae Pseudomonadaceae Summary of strains isolated belonging to the class Gammaproteobacteria and their taxonomic affiliations P hylum Table 1. Table Gammaproteobacteria February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 27

(1) (2) (3) (4) (5) (6)

(7) (8) (9) (10) (11) (12)

(13) (14) (15) (16) (17) (18)

(19) (20) (21) (22) (23) (24)

(25) (26) (27) (28) (29) (30)

(31) (32) (33) (34) (35) (36)

(37) (38) (39)

Fig. 1. Transmission electron micrographs or scanning electron micrographs of cells of the strains isolated in the study. Strains: 1, 4003; 2, 4231; 3, 7026; 4, 7034; 5, 9023; 6, BE6-1; 7, CAU 1046; 8, CAU 1335; 9, DS041; 10, EC2D12; 11, GM22; 12, Gsoil 3009; 13, HMF7415; 14, HMF8002; 15, HMF8004; 16, HMF8044; 17, IMCC25635; 18, IMCC25639; 19, IMCC25642; 20, IMCC25643; 21, IMCC25647; 22, IMCC25652; 23, KB012; 24, LGGG06; 25, LPB0132; 26, LPB0153; 27, LPB0156; 28, LPB0159; 29, POB9; 30, SFD22; 31, SN5; 32, SN10; 33, UL158; 34, UL222; 35, UL498-1; 36, UL506; 37, YC7-24; 38, YHY1; 39, ZOD21. 28 Journal of Species Research Vol. 7, No. 1

Fig. 2. Neighbor-joining phylogenetic tree, based on 16S rRNA gene sequences, showing the relationships between the strains isolated in this study and their relatives of the order Aeromonadales and Alteromonadales in the class Gammaproteobacteria. Bootstrap values (>50%) are shown at branching points. Filled circles indicate that the corresponding nodes were also recovered in the trees generated with the maximum-likelihood and maximum-parsimony algorithms, while open circles indicate that the corresponding nodes were also recovered in the tree generated with one of these algorithms. Bar, 0.02 substitutions per nucleotide position.

Fig. 3. Neighbor-joining phylogenetic tree, based on 16S rRNA gene sequences, showing the relationships between the strains isolated in this study and their relatives of the order Arenicellales, Vibrionales and Xanthomonadales in the class Gammaproteobacteria. Bootstrap values (>50%) are shown at branching points. Filled circles indicate that the corresponding nodes were also recovered in the trees generated with the maximum-likelihood and maximum-parsimony algorithms, while open circles indicate that the corresponding nodes were also recovered in the tree generated with one of these algorithms. Bar, 0.02 substitutions per nucleotide position. smooth, raised and white colored after incubation for but not D-maltose, capric acid, adipic acid and pheny- 2 days on R2A at 30℃. Positive for nitrate reduction, lacetic acid. Strain 7026 (=NIBRBAC000498574) was indole production, glucose fermentation, esculin hydro- isolated from a sediment sample, Seoul, Korea. lysis, gelatin hydrolysis and β-galactosidase, but negative for arginine dihydrolase and urease in API 20NE. Description of Yokenella regensburgei 7034 D-glucose, L-arabinose, D-mannose, D-mannitol, N- acetyl-glucosamine, potassium gluconate, malic acid Cells are Gram-stain-negative, flagellated, non-pig- and trisodium citrate are utilized as sole carbon sources, mented and rod-shaped. Colonies are circular, entire, February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 29

Fig. 4. Neighbor-joining phylogenetic tree, based on 16S rRNA gene sequences, showing the relationships between the strains isolated in this study and their relatives of the order Enterobacteriales in the class Gammaproteobacteria. Bootstrap values (>50%) are shown at branching points. Filled circles indicate that the corresponding nodes were also recovered in the trees generated with the maximum-likelihood and maximum-parsimony algorithms, while open circles indicate that the corresponding nodes were also recovered in the tree generated with one of these algorithms. Bar, 0.02 substitutions per nucleotide position.

Fig. 5. Neighbor-joining phylogenetic tree, based on 16S rRNA gene sequences, showing the relationships between the strains isolated in this study and their relatives of the order Pseudomonadales in the class Gammaproteobacteria. Bootstrap values (>50%) are shown at branching points. Filled circles indicate that the corresponding nodes were also recovered in the trees generated with the maximum-likelihood and maximum-parsimony algorithms, while open circles indicate that the corresponding nodes were also recovered in the tree generated with one of these algorithms. Bar, 0.02 substitutions per nucleotide position. smooth, raised and white colored after incubation for Description of Dickeya chrysanthemi 9023 2 days on R2A at 30℃. Positive for nitrate reduction, indole production, glucose fermentation, esculin hydro- Cells are Gram-stain-negative, non-flagellated, non- lysis, gelatin hydrolysis and β-galactosidase, but nega- pigmented and rod-shaped. Colonies are irregular, un- tive for arginine dihydrolase and urease in API 20NE. dulate, glistening and white colored after incubation for D-glucose, L-arabinose, D-mannose, D-mannitol, N- 2 days on R2A at 30℃. Positive for nitrate reduction, acetyl-glucosamine, potassium gluconate, malic acid indole production, glucose fermentation, esculin hydro- and trisodium citrate are utilized as sole carbon sources, lysis, gelatin hydrolysis and β-galactosidase, but nega- but not D-maltose, capric acid, adipic acid and pheny- tive for arginine dihydrolase and urease in API 20NE. lacetic acid. Strain 7034 (=NIBRBAC000498575) was D-glucose, L-arabinose, D-mannose, D-mannitol, N- isolated from a sediment sample, Seoul, Korea. acetyl-glucosamine, D-maltose, potassium gluconate, 30 Journal of Species Research Vol. 7, No. 1 malic acid and trisodium citrate are utilized as sole car- pigmented and rod-shaped. Colonies are circular, convex bon sources, but not capric acid, adipic acid and pheny- and pale yellow colored after 4 days of incubation on lacetic acid. Strain 9023 (=NIBRBAC000498576) was R2A at 25°C. Positive for nitrate reduction, but negative isolated from a sediment sample, Seoul, Korea. for indole production, glucose fermentation, arginine dihydrolase, urease, esculin hydrolysis, gelatin hydrolysis Description of Escherichia fergusonii BE6-1 and β-galactosidase in API 20NE. D-glucose, L-arabinose, capric acid, malic acid and trisodium citrate are Cells are Gram-stain-negative, non-flagellated, non- utilized as sole carbon sources, but not D-mannose, pigmented and rod-shaped. Colonies are circular, con- D-mannitol, N-acetyl-glucosamine, D-maltose, potassi- vex, entire and cream colored after incubation for 4 days um gluconate, adipic acid and phenylacetic acid. Strain on TSA at 30°C. Positive for nitrate reduction, glucose DS041 ( =NIBRBAC000498630) was isolated from a fermentation, esculin hydrolysis and β-galactosidase, freshwater sample, Seoul, Korea. but negative for indole production, arginine dihydrolase, urease and gelatin hydrolysis in API 20NE. D-glucose, L-arabinose, D-mannose, D-mannitol, N-acetyl-glucos- Description of Pseudoalteromonas lipolytica EC2D12 amine, D-maltose, potassium gluconate, malic acid and Cells are Gram-stain-negative, non-flagellated, non- phenylacetic acid are utilized as sole carbon sources, but pigmented and rod-shaped. Colonies are circular, trans- not capric acid, adipic acid and trisodium citrate. Strain parent, smooth, umbonate and cream colored after in-

BE6-1 ( =NIBRBAC000498660) was isolated from a cubation for 3 days on MA at 25℃. Positive for nitrate tidal flat sample, Eulsuk Island, Korea. reduction, esculin hydrolysis and gelatin hydrolysis, but negative for indole production, glucose fermentation, Description of Microbulbifer taiwanensis CAU 1046 arginine dihydrolase, urease and β-galactosidase in API Cells are Gram-stain-negative, non-flagellated, non- 20NE. Does not utilize D-glucose, L-arabinose, D-man- pigmented and rod-shaped. Colonies are circular, glis- nose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, capric acid, adipic acid, malic acid, tening, convex and pale yellow colored after incubation for 3 days on MA at 30℃. Positive for esculin hydroly- trisodium citrate and phenylacetic acid. Strain EC2D12 = sis and gelatin hydrolysis, but negative for nitrate reduc- ( NIBRBAC000498484) was isolated from an Ecklo- tion, indole production, glucose fermentation, arginine nia cava sample, Yeosu, Korea. dihydrolase, urease and β-galactosidase in API 20NE. L-arabinose and D-maltose are utilized as sole carbon Description of Serratia quinivorans GM22 sources, but not D-glucose, D-mannose, N-acetyl-glu- Cells are Gram-stain-negative, non-flagellated, non- cosamine, potassium gluconate, capric acid, adipic acid, pigmented and rod-shaped. Colonies are circular, malic acid, trisodium citrate and phenylacetic acid. opaque, smooth, convex and cream colored after in- = Strain CAU 1046 ( NIBRBAC000498497) was isolat- cubation for 1 day on LB at 25℃. Positive for nitrate ed from a soil sample, Jeju Island, Korea. reduction, glucose fermentation, esculin hydrolysis, gelatin hydrolysis and β-galactosidase, but negative for in- Description of Pseudomonas punonensis CAU 1335 dole production, arginine dihydrolase and urease in API Cells are Gram-stain-negative, flagellated, non-pig- 20NE. D-glucose, L-arabinose, D-mannose, D-manni- mented and rod-shaped. Colonies are small circular, tol, N-acetyl-glucosamine, D-maltose, potassium gluco- convex and cream colored after incubation for 3 days on nate, capric acid, malic acid and trisodium citrate are uti- pH 8 MA at 30℃. Positive for nitrate reduction, glucose lized as sole carbon sources, but not adipic acid and phe- fermentation, arginine dihydrolase, urease and esculin nylacetic acid. Strain GM22 ( =NIBRBAC000498486) hydrolysis, but negative for indole production, gelatin was isolated from a Tenebrio molitor sample, Gokseong, hydrolysis and β-galactosidase in API 20NE. D-glu- Korea. cose, L-arabinose, N-acetylglucosamine, D-maltose, capric acid, malic acid and trisodium citrate are utilized Description of Rhodanobacter spathiphylli Gsoil 3009 as sole carbon sources, but not D-mannose, D-mannitol, Cells are Gram-stain-negative, non-flagellated, non- potassium gluconate, adipic acid and phenylacetic acid. pigmented and rod-shaped. Colonies are circular, con- Strain CAU 1335 (=NIBRBAC000498504) was isolat- vex, glistening and yellow colored after incubation for 3 ed from a seawater sample, Jeju Island, Korea. days on R2A at 25℃. Positive for nitrate reduction and esculin hydrolysis, but negative for indole production, Description of Acinetobacter junii DS041 glucose fermentation, arginine dihydrolase, urease, gela- Cells are Gram-stain-negative, non-flagellated, non- tin hydrolysis and β-galactosidase in API 20NE. D-glu- February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 31 cose, D-mannose, N-acetyl-glucosamine and D-maltose Description of Salinivibrio sharmensis HMF8044 are utilized as sole carbon sources, but not L-arabinose, Cells are Gram-stain-negative, non-flagellated, non- D-mannitol, potassium gluconate, capric acid, adipic pigmented and curved-rod-shaped. Colonies are circular, acid, malic acid, trisodium citrate and phenylacetic acid. convex, entire and beige colored after incubation for 2 Strain Gsoil 3009 (=NIBRBAC000498590) was isolat- days on MA at 37℃. Positive for esculin hydrolysis, but ed from a ginseng field soil sample, Anseong, Korea. negative for nitrate reduction, indole production, glucose fermentation, arginine dihydrolase, urease, gelatin hydro- Description of Erwinia billingiae HMF7415 lysis and β-galactosidase in API 20NE. Dose not utilize D- Cells are Gram-stain-negative, non-flagellated, non- glucose, L-arabinose, D-mannose, D-mannitol, N-acetyl- pigmented and rod to oval-shaped. Colonies are circu- glucosamine, D-maltose, potassium gluconate, capric lar, convex, entire and beige colored after incubation for acid, adipic acid, malic acid, trisodium citrate and phenyl

3 days on R2A at 25℃. Positive for nitrate reduction, acetic acid. Strain HMF8044 (=NIBRBAC000498458) glucose fermentation, esculin hydrolysis and β-galac- was isolated from a saltern sample, Sinan, Korea. tosidase, but negative for indole production, arginine dihydrolase, urease and gelatin hydrolysis in API 20NE. Description of Shewanella algae IMCC25635 D-glucose, L-arabinose, D-mannose, D-mannitol, N- Cells are Gram-stain-negative, flagellated, non-pig- acetyl-glucosamine, D-maltose, potassium gluconate mented and rod-shaped. Colonies are circular, entire, and malic acid are utilized as sole carbon sources, but convex and orange colored after incubation for 3 days not capric acid, adipic acid, trisodium citrate and pheny- on MA at 20℃. Positive for indole production, gelatin = lacetic acid. Strain HMF7415 ( NIBRBAC000498453) hydrolysis and β-galactosidase, but negative for nitrate was isolated from a tree sample, Yongin, Korea. reduction, glucose fermentation, arginine dihydrolase, urease and esculin hydrolysis in API 20NE. D-maltose Description of Salinivibrio costicola subsp. and trisodium citrate are utilized as sole carbon sources, vallismortis HMF8002 but not D-glucose, L-arabinose, D-mannose, D-manni- Cells are Gram-stain-negative, non-flagellated, non- tol, N-acetyl-glucosamine, potassium gluconate, capric pigmented and curved-rod-shaped. Colonies are circu- acid, adipic acid, malic acid and phenylacetic acid. = lar, convex, entire and red colored after incubation for 2 Strain IMCC25635 ( NIBRBAC000498534) was iso- days on MA at 37℃. Positive for glucose fermentation, lated from a tidal flat sample, Mo Island, Korea. arginine dihydrolase and gelatin hydrolysis, but negative for nitrate reduction, indole production, urease, esculin Description of Marinobacterium nitratireducens hydrolysis and β-galactosidase in API 20NE. D-glu- IMCC25639 cose, D-mannose, N-acetyl-glucosamine, potassium glu- Cells are Gram-stain-negative, non-flagellated, non- conate and malic acid are utilized as sole carbon sourc- pigmented and rod-shaped. Colonies are circular, entire, es, but not L-arabinose, D-mannitol, D-maltose, capric convex and yellow colored after incubation for 3 days on acid, adipic acid, trisodium citrate and phenylacetic acid. MA at 20℃. Negative for nitrate reduction, indole pro-

Strain HMF8002 ( =NIBRBAC000498456) was isolat- duction, glucose fermentation, arginine dihydrolase, ure- ed from a saltern sample, Sinan, Korea. ase, esculin hydrolysis, gelatin hydrolysis and β-galactosidase in API 20NE. Does not utilize D-glucose, L-arab- Description of Vibrio ruber HMF8004 inose, D-mannose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, capric acid, adipic Cells are Gram-stain-negative, non-flagellated, non- acid, malic acid, trisodium citrate and phenylacetic acid. pigmented and curved-rod-shaped. Colonies are circu- Strain IMCC25639 ( =NIBRBAC000498538) was iso- lar, convex, entire and red colored after incubation for lated from a tidal flat sample, Mo Island, Korea. 2 days on MA at 30℃. Positive for nitrate reduction, glucose fermentation, esculin hydrolysis, gelatin hydro- Description of Microbulbifer mangrove IMCC25642 lysis and β-galactosidase, but negative for indole production, arginine dihydrolase and urease in API 20NE. Cells are Gram-stain-negative, non-flagellated, non- D-glucose, L-arabinose, D-mannose, D-mannitol and pigmented and coccoid-shaped. Colonies are circular, D-maltose are utilized as sole carbon sources, but not convex, entire and yellow colored after incubation for N-acetyl-glucosamine, potassium gluconate, capric acid, 3 days on MA at 20℃. Positive for nitrate reduction, adipic acid, malic acid, trisodium citrate and pheny- gelatin hydrolysis and β-galactosidase, but negative for lacetic acid. Strain HMF8004 (=NIBRBAC000498457) indole production, glucose fermentation, arginine di- was isolated from a saltern sample, Sinan, Korea. hydrolase, urease and esculin hydrolysis in API 20NE. 32 Journal of Species Research Vol. 7, No. 1

Does not utilize D-glucose, L-arabinose, D-mannose, lated from a plant root sample, Jo Island, Korea. D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, capric acid, adipic acid, malic acid, triso- Description of Aeromonas allosaccharophila KB012 dium citrate and phenylacetic acid. Strain IMCC25642 Cells are Gram-stain-negative, non-flagellated, non- (=NIBRBAC000498541) was isolated from a tidal flat pigmented and oval-shaped. Colonies are circular, con- sample, Mo Island, Korea. vex, smooth, opaque and white colored after incubation for 4 days on R2A at 25℃. Positive for nitrate reduc- Description of Marinobacter hydrocarbonoclasticus tion, indole production, glucose fermentation, arginine IMCC25643 dihydrolase, esculin hydrolysis, gelatin hydrolysis and Cells are Gram-stain-negative, non-flagellated, non- β-galactosidase, but negative for urease in API 20NE. pigmented and rod-shaped. Colonies are circular, con- D-glucose, L-arabinose, D-mannose, D-mannitol, vex, entire and white colored after incubation for 3 N-acetyl-glucosamine, D-maltose, potassium gluconate, days on MA at 20℃. Positive for nitrate reduction, but capric acid, malic acid and trisodium citrate are utilized negative for indole production, glucose fermentation, as sole carbon sources, but not adipic acid and pheny- arginine dihydrolase, urease, esculin hydrolysis, gelatin lacetic acid. Strain KB012 ( =NIBRBAC000498636) hydrolysis and β-galactosidase in API 20NE. Does not was isolated from a freshwater sample, Gunsan, Korea utilize D-glucose, L-arabinose, D-mannose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, Description of Psychrobacter marincola LGGG06 capric acid, adipic acid, malic acid, trisodium citrate Cells are Gram-stain-negative, non-flagellated, non- and phenylacetic acid. Strain IMCC25643 ( =NIBR pigmented and oval-shaped. Colonies are circular, BAC000498542) was isolated from a tidal flat sample, slightly convex and pale yellow colored after incubation Mo Island, Korea. for 3 days on R2A at 25℃. Positive for nitrate reduction and glucose fermentation, but negative for indole pro- Description of Acinetobacter radioresistens duction, arginine dihydrolase, urease, esculin hydroly- IMCC25647 sis, gelatin hydrolysis and β-galactosidase in API 20NE. Does not utilize D-glucose, L-arabinose, D-mannose, D- Cells are Gram-stain-negative, non-flagellated, non- mannitol, N-acetyl-glucosamine, D-maltose, potassium pigmented and rod-shaped. Colonies are circular, con- gluconate, capric acid, adipic acid, malic acid, trisodium vex, entire and white colored after incubation for 3 days citrate and phenylacetic acid. Strain LGGG06 ( =NI- on MA at 20℃. Negative for nitrate reduction, indole BRBAC000498632) was isolated from a seawater sam- production, glucose fermentation, arginine dihydrolase, ple, Jeju Island, Korea. urease, esculin hydrolysis, gelatin hydrolysis and β-galactosidase in API 20NE. L-arabinose, adipic acid and Description of Marinobacter oulmenensis LPB0132 malic acid are utilized as sole carbon sources, but not Cells are Gram-stain-negative, flagellated, non-pig- D-glucose, D-mannose, D-mannitol, N-acetyl-glucos- mented and rod-shaped. Colonies are circular, entire, amine, D-maltose, potassium gluconate, capric acid, tri- convex and beige colored after incubation for 1 day on sodium citrate and phenylacetic acid. Strain IMCC25647 MA at 25℃. Positive for nitrate reduction and gelatin ( =NIBRBAC000498546) was isolated from a plant hydrolysis, but negative for indole production, glu- root sample, Jo Island, Korea. cose fermentation, arginine dihydrolase, urease, esculin hydrolysis and β-galactosidase in API 20NE. N-ace- Description of Pseudomonas alcaliphila IMCC25652 tyl-glucosamine and malic acid are utilized as sole carbon sources, but not D-glucose, L-arabinose, D-man- Cells are Gram-stain-negative, flagellated, non-pig- nose, D-mannitol, D-maltose, potassium gluconate, mented and rod-shaped. Colonies are circular, convex, capric acid, adipic acid, trisodium citrate and pheny- entire and white colored after incubation for 3 days on lacetic acid. Strain LPB0132 (=NIBRBAC000498520) MA at 20℃. Positive for esculin hydrolysis, but neg- was isolated from an Urechis unicinctus sample, Jumun- ative for nitrate reduction, indole production, glucose jin, Korea. fermentation, arginine dihydrolase, urease, gelatin hydrolysis and β-galactosidase in API 20NE. L-arabinose, Description of Shewanella algidipiscicola LPB0153 N-acetyl-glucosamine, capric acid, adipic acid, trisodium citrate and phenylacetic acid are utilized as sole Cells are Gram-stain-negative, flagellated, non-pig- carbon sources, but not D-glucose, D-mannose, D-man- mented and rod-shaped. Colonies are circular, entire, nitol, D-maltose, potassium gluconate and malic acid. convex and beige colored after incubation for 1 day on Strain IMCC25652 ( =NIBRBAC000498551) was iso- MA at 25℃. Positive for nitrate reduction and esculin February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 33 hydrolysis, but negative for indole production, glucose from a wastewater sample, Gwangyang, Korea. fermentation, arginine dihydrolase, urease, gelatin hydrolysis and β-galactosidase in API 20NE. N-Acetyl- Description of Vibrio fortis SFD22 glucosamine and malic acid are utilized as sole carbon Cells are Gram-stain-negative, non-flagellated, non- sources, but not D-glucose, L-arabinose, D-mannose, pigmented and rod-shaped. Colonies are opaque, D-mannitol, D-maltose, potassium gluconate, capric smooth, low convex and cream colored after incubation acid, adipic acid, trisodium citrate and phenylacetic acid. for 3 days on MA at 25℃. Positive for esculin hydroly-

Strain LPB0153 (=NIBRBAC000498525) was isolated sis, but negative for nitrate reduction, indole production, from a tidal flat sample, Jebu Island, Korea. glucose fermentation, arginine dihydrolase, urease, gelatin hydrolysis and β-galactosidase in API 20NE. Dose Description of Marinobacter sediminum LPB0156 not utilize D-glucose, L-arabinose, D-mannose, D-man- Cells are Gram-stain-negative, flagellated, non-pig- nitol, N-acetyl-glucosamine, D-maltose, potassium glu- mented and rod-shaped. Colonies are circular, entire, conate, capric acid, adipic acid, malic acid, trisodium convex and yellow colored after 1 day of incubation citrate and phenylacetic acid. Strain SFD22 ( =NIBR on MA at 25℃. Positive for nitrate reduction, but neg- BAC000498472) was isolated from a Sargassum fulvel- ative for indole production, glucose fermentation, ar- lum sample, Jeju Island, Korea. ginine dihydrolase, urease, esculin hydrolysis, gelatin hydrolysis and β-galactosidase in API 20NE. Does not Description of Xanthomonas saccahri SN5 utilize D-glucose, L-arabinose, D-mannose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium glu- Cells are Gram-stain-negative, non-flagellated, non- conate, capric acid, adipic acid, malic acid, trisodium pigmented and rod-shaped. Colonies are circular, raise, citrate and phenylacetic acid. Strain LPB0156 (=NIBR entire and pale-yellow colored after incubation for 2 BAC000498527) was isolated from a tidal flat sample, days on MA at 30℃. Positive for esculin hydrolysis, Jebu Island, Korea. gelatin hydrolysis and β-galactosidase, but negative for nitrate reduction, indole production, glucose fermen- Description of Marinobacter guineae LPB0159 tation, arginine dihydrolase and urease in API 20NE. D-glucose, L-arabinose, D-mannose, N-acetyl-glucos- Cells are Gram-stain-negative, flagellated, non-pig- amine, D-maltose, malic acid and trisodium citrate are mented and rod-shaped. Colonies are circular, entire, utilized as sole carbon sources, but not D-mannitol, po- convex and beige colored after 1 day of incubation on tassium gluconate, capric acid, adipic acid and pheny- MA at 25℃. Positive for nitrate reduction, but nega- lacetic acid. Strain SN5 ( =NIBRBAC000498425) was tive for indole production, glucose fermentation, ar- isolated from a sewage treatment plant sample, Busan, ginine dihydrolase, urease, esculin hydrolysis, gelatin Korea. hydrolysis and β-galactosidase in API 20NE. Does not utilize D-glucose, L-arabinose, D-mannose, D-manni- Description of Pantoea stewartii subsp. indologenes tol, N-acetyl-glucosamine, D-maltose, potassium glu- SN10 conate, capric acid, adipic acid, malic acid, trisodium citrate and phenylacetic acid. Strain LPB0159 (=NIBR Cells are Gram-stain-negative, non-flagellated, non- BAC000498530) was isolated from a tidal flat sample, pigmented and rod-shaped. Colonies are circular, raise, Jebu Island, Korea. entire and pale yellow colored after incubation for 2 days on MA at 30℃. Positive for esculin hydrolysis, Description of Psychrobacter faecalis POB9 gelatin hydrolysis and β-galactosidase, but negative for nitrate reduction, indole production, glucose fermen- Cells are Gram-stain-negative, non-flagellated, non- tation, arginine dihydrolase and urease in API 20NE. pigmented and coccobacilli-shaped. Colonies are D-glucose, L-arabinose, D-mannose, N-acetyl-glucos- opaque, circular, smooth, convex and cream colored af- amine, D-maltose, malic acid and trisodium citrate are ter incubation for 2 days on R2A at 25℃. Positive for utilized as sole carbon sources, but not D-mannitol, po- nitrate reduction and glucose fermentation, but nega- tassium gluconate, capric acid, adipic acid and pheny- tive for indole production, arginine dihydrolase, urease, lacetic acid. Strain SN10 (=NIBRBAC000498426) was esculin hydrolysis, gelatin hydrolysis and β-galactosi- isolated from a sewage treatment plant sample, Busan, dase in API 20NE. Dose not utilize D-glucose, L-arab- Korea. inose, D-mannose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, capric acid, adipic Description of Psychrobacter piscatorii UL158 acid, malic acid, trisodium citrate and phenylacetic acid. Strain POB9 ( =NIBRBAC000498473) was isolated Cells are Gram-stain-negative, non-flagellated, non- 34 Journal of Species Research Vol. 7, No. 1 pigmented and rod-shaped. Colonies are circular, β-galactosidase in API 20NE. Does not utilize D-glu- opaque, convex and beige colored after incubation for cose, L-arabinose, D-mannose, D-mannitol, N-ace- 3 days on TSA at 25°C. Positive for esculin hydrolysis, tyl-glucosamine, D-maltose, potassium gluconate, capric but negative for nitrate reduction, indole production, acid, adipic acid, malic acid, trisodium citrate and phe- glucose fermentation, arginine dihydrolase, urease, nylacetic acid. Strain UL506 (=NIBRBAC000498619) gelatin hydrolysis and β-galactosidase in API 20NE. was isolated from a soil sample, Ulleung Island, Korea. L-arabinose and malic acid are utilized as sole carbon sources, but not D-glucose, D-mannose, D-mannitol, Description of Pseudomonas cuatrocienegasensis N-acetyl-glucosamine, D-maltose, potassium gluconate, YC7-24 capric acid, adipic acid, trisodium citrate and pheny- Cells are Gram-stain-negative, flagellated, non-pig- lacetic acid. Strain UL158 ( =NIBRBAC000498615) mented and rod-shaped. Colonies are circular, convex, was isolated from a seawater sample, Ulleung Island, entire and light yellow colored after incubation for 2 Korea. days on TSA at 30°C. Positive for nitrate reduction, but negative for indole production, glucose fermentation, Description of Haemophilus piscium UL222 arginine dihydrolase, urease, esculin hydrolysis, gelatin Cells are Gram-stain-negative, non-flagellated, non- hydrolysis and β-galactosidase in API 20NE. D-glu- pigmented and rod-shaped. Colonies are circular, cose, potassium gluconate, malic acid and trisodium ci- opaque, convex and beige colored after incubation for trate are utilized as sole carbon sources, but not L-arab- 3 days on TSA at 25℃. Positive for nitrate reduction, inose, D-mannose, D-mannitol, N-acetyl-glucosamine, indole production, glucose fermentation, esculin hydro- D-maltose, capric acid, adipic acid and phenylacetic lysis, gelatin hydrolysis and β-galactosidase, but nega- acid. Strain YC7-24 (=NIBRBAC000498655) was iso- tive for arginine dihydrolase and urease in API 20NE. lated from a cave sample, Jeju Island, Korea. D-glucose, L-arabinose, D-mannose, D-mannitol, N-acetyl-glucosamine, D-maltose, potassium gluconate, Description of Rhodanobacter glycinis YHY1 capric acid, malic acid and trisodium citrate are utilized Cells are Gram-stain-negative, flagellated, non-pig- as sole carbon sources, but not adipic acid and pheny- mented and rod-shaped. Colonies are circular, smooth, lacetic acid. Strain UL222 ( =NIBRBAC000498616) convex and ivory colored after incubation for 2 days was isolated from a seawater sample, Ulleung Island, on R2A at 25℃. Negative for nitrate reduction, indole Korea. production, glucose fermentation, arginine dihydrolase, urease, esculin hydrolysis, gelatin hydrolysis and β-ga- Description of Pseudomonas guineae UL498-1 lactosidase in API 20NE. N-acetyl-glucosamine and Cells are Gram-stain-negative, flagellated, pigmented D-maltose are utilized as sole carbon sources, but not and rod-shaped. Colonies are circular, convex, smooth, D-glucose, L-arabinose, D-mannose, D-mannitol, po- glistening and pale yellow colored after incubation for 2 tassium gluconate, capric acid, adipic acid, malic acid, days on TSA at 25°C. Positive for gelatin hydrolysis, but trisodium citrate and phenylacetic acid. Strain YHY1 negative for nitrate reduction, indole production, glu- ( =NIBRBAC000498406) was isolated from a sewage cose fermentation, arginine dihydrolase, urease, esculin treatment plant sample, Busan, Korea. hydrolysis, β-galactosidase in API 20NE. D-glucose, capric acid, malic acid and trisodium citrate are utilized Description of Arenicella chitinivorans ZOD21 as sole carbon sources, but not L-arabinose, D-mannose, Cells are Gram-stain-negative, non-flagellated, non- D-mannitol, N-acetyl-glucosamine, D-maltose, potassi- pigmented and rod-shaped. Colonies are irregular, circu- um gluconate, adipic acid and phenylacetic acid. Strain lar, smooth, entire and yellow colored after incubation UL498-1 (=NIBRBAC000498618) was isolated from a for 5 days on MA at 25℃. Positive for esculin hydroly- seawater sample, Ulleung Island, Korea. sis and gelatin hydrolysis, but negative for nitrate reduction, indole production, glucose fermentation, arginine Description of Pseudomonas salina UL506 dihydrolase, urease and β-galactosidase in API 20NE. Cells are Gram-stain-negative, flagellated, non-pig- Dose not utilize D-glucose, L-arabinose, D-mannose, mented and rod-shaped. Colonies are circular, convex, D-mannitol, N-acetyl-glucosamine, D-maltose, potas- entire and pale yellow colored after incubation for 3 sium gluconate, capric acid, adipic acid, malic acid, days on TSA at 30°C. Negative for nitrate reduction, in- trisodium citrate and phenylacetic acid. Strain ZOD21 dole production, glucose fermentation, arginine dihydro- ( =NIBRBAC000498478) was isolated from a Zostera lase, urease, esculin hydrolysis, gelatin hydrolysis and marina sample, Seosan, Korea. February 2018 Kim et al. - Unrecorded species of Gammaproteobacteria 35

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