A Report of 35 Unreported Bacterial Species in Korea, Belonging to the Phylum Firmicutes

Home , Staphylococcus argenteus, Staphylococcus lentus

Journal of Species Research 8(4):337-350, 2019

A report of 35 unreported bacterial species in Korea, belonging to the phylum Firmicutes

Min-gyung Baek1, Wonyong Kim3, Chang-Jun Cha4, Kiseong Joh5, Seung-Bum Kim6, Myung Kyum Kim7, Chi-Nam Seong8 and Hana Yi1,2,*

1Department of Public Health Science, Korea University, Seoul 02841, Republic of Korea 2School of Biosystem and Biomedical Science, Department of Public Health Science, Korea University, Seoul 02841, Republic of Korea 3Department of Microbiology, Chung-Ang University College of Medicine, Seoul 06974, Republic of Korea 4Department of Biotechnology, Chung-Ang University, Anseong 17546, Republic of Korea 5Department of Bioscience and Biotechnology, Hankuk University of Foreign Studies, Geonggi 02450, Republic of Korea 6Department of Microbiology, Chungnam National University, Daejeon 34134, Republic of Korea 7Department of Bio and Environmental Technology, College of Natural Science, Seoul Women’s University, Seoul 01797, Republic of Korea 8Department of Biology, Sunchon National University, Suncheon 57922, Republic of Korea

*Correspondent: [email protected]

In an investigation of indigenous prokaryotic species in Korea, a total of 35 bacterial strains assigned to the phylum Firmicutes were isolated from diverse habitats including natural and artificial environments. Based on their high 16S rRNA gene sequence similarity (>98.7%) and formation of robust phylogenetic clades with species of validly published names, the isolates were identified as 35 species belonging to the ordersBacillales (the family Bacillaceae, Paenibacillaceae, Planococcaceae, and Staphylococcaceae) and Lactobacillales (Aerococcaceae, Enterococcaceae, Lactobacillaceae, Leuconostocaceae, and Streptococcaceae). Since these 35 species in Korean environments has not been reported in any official report, we identified them as unrecorded bacterial species and investigated them taxonomically. The newly found unrecorded species belong to 20 species in the order Bacillales and 15 species in the order Lactobacillales. The morphological, cultural, physiological, and biochemical properties of the isolates were examined and the descriptive information of the 35 previously unrecorded species is provided here. Keywords: ‌16S rRNA, Firmicutes, unrecorded species

Introduction tain teichoic acid instead (De Vos et al., 2011). Respira- tion is aerobic, anaerobic, or facultative anaerobic. Cells The phylum Firmicutes (Gibbons and Murray, 1978) is are spherical-, straight-, curved-, helical-, or filamen- a large group consisting of 7 classes (Bacilli, Clostridia, tous-shaped. Most members are chemoorganotrophs, but Erysipelotrichia, Limnochordia, Negativicutes, Thermo- a few are anoxygenic photoheterotrophs (De Vos et al., lithobacteria, and Tissierellia), and at least 39 families 2011). and 274 genera. The taxonomy of this phylum is the The orders Bacillales and Lactobacillales are the larg- subject of numerous recent rearrangements because it is est and most popular members of the Firmicutes. The phenotypically and phylogenetically diverse. Traditional- order Bacillales is generally characterized by the forma- ly Firmicutes has been known as gram-positive with low tion of endospores and possession of menaquinone 7, and G+C (mol% G+C content of DNA is <50), but some contains 9 familes named Bacillaceae, Alicyclobacillace- gram-negative members are also present in the phylum ae, Listeriaceae, Paenibacillaceae, Pasteuriaceae, Pla- (e.g., Vellonellaceae, Syntrophomonadaceae). The cell nococcaceae, Sporolactobacillaceae, Staphylococcaceae, walls are rigid by containing muramic acid, but some con and Thermoactinomycetaceae (De Vos et al., 2011). The 338 Journal of Species Research Vol. 8, No. 4 order Lactobacillales is generally characterized by being were compared with sequences of validly published type facultative anaerobic respiration without forming an en- strains using the EzBioCloud server (Yoon et al., 2017). dospore, and contains 7 families named Lactobacillaceae, Based on pairwise sequence similarity, an isolate showing Aerococcaceae, Carnobacteriaceae, Enterococcaceae, 98.7% or higher similarity to any type strain of the phy- Leuconostocaceae, Streptococcaceae, and Clostridiacea lum Firmicutes, but whose presence has not been report- (De Vos et al., 2011). ed in Korea, was identified as an unrecorded Firmicutes According to the recent microbiome studies, Firmicutes species. The sequences were aligned using EzEditor (Jeon has a significant impact on human health and disease, et al., 2014) and phylogenetic trees were reconstructed such as obesity, autoimmune disease, allergic diseases, using maximum-likelihood and neighbor-joining methods and intestinal disease. For example, in irritable bowel implemented in MEGA7 (Kumar et al., 2016). syndrome, the increased Firmicutes abundance was pos- Cellular morphology and cell size were examined by itively correlated with the disease severity (Rajilić-Sto- either transmission electron microscopy or scanning elec- janović et al., 2011; Jeffery et al., 2012). In obesity, the tron microscopy. Gram staining was performed using abundance of Firmicutes increases proportionally because a Gram-staining kit. Biochemical characteristics were the bacteria leads to decrease in overall metabolic diver- tested by using API 20NE or API 20A kits (bioMérieux) sity in the human gut (Bäckhed et al., 2004; Ley et al., according to the manufacturer’s instructions. Oxidase ac- 2005; Turnbaugh et al., 2009). tivity was evaluated using 1% tetramethyl-p-phenylenedi- During the indigenous bacterial diversity survey per- amine (bioMérieux). formed by National Institute of Biological Resources in Korea, numerous bacterial strains belonging to the phylum Firmicutes were isolated from various sources Results and Discussion including natural and artificial environments, as well as animals and human. As a result of preliminary phyloge- Thirty-five Firmicutes strains were isolated from vari- netic analysis using partial 16S rRNA gene sequences, 35 ous environmental habitats and identified as unrecorded of them were identified to belong to the orders Bacillales bacterial species by having 16s rRNA gene sequence sim- and Lactobacillales with a possibility to be recognized as ilarity 98.7% or higher but never been reported in Korea. unrecorded bacterial species. Thus, the taxonomy of the The similarity-based identification was also supported by new isolates was investigated and the presence of 35 in- phylogenetic trees (Fig. 1). Each isolate formed a mono- digenous species in Korea is reported here. phyletic clade with the type strain of identified bacterial species, confirming the proper assignment of the isolate to the species with published names. The tree topologies Materials and Methods of neighbor-joining and maximum likelihood trees were almost identical showing the close relationship of the iso- As sources of bacteria, samples of lichen, sea salt, sea lates and valid type strains of the family Firmicutes. The water, sea sand, soil, moss, mouse gut, mine drainage, hu- taxonomic affiliations of the isolates are summarized in man sputum, human feces, and wood were collected from Table 1. natural and artificial environments in Republic of Korea The 35 isolates belonged to 2 orders of the Firmicutes; (Table 1). Bacterial culture media were used for standard 20 isolates to the order Bacillales, 15 isolates to the or- dilution plating technique, including Reasoner’s 2A (R2A; der Lactobacillales. In the order Bacillales, the 20 iso- BD), Marine Agar 2216 (MA; BD), Mueller Hinton Agar lates further grouped into 4 bacterial families; 5 isolates (MH; BD), Luria-Bertani Agar (LB; BD), Glucose Yeast (HMF5940, GA079, 18JY44-4, LPB0245, and CAU Extract Agar (GYE; BD), Anaerobe Basal Agar (AB; Ox- 1495) of the family Bacillaceae; 8 isolates (18S2L7, CAU oid), Tryptic Soy Yeast Agar (TSY; BD), and Lactobacilli 1503, HMF7158, 18H7A6, MMS18-CY120, HMF7562, MRS Agar (MRS; BD). The spread plates were incubated GY1, and 18JY2-4) of the family Paenibacillaceae; 5 at 25-37℃ for 1-7 days and subjected to single colony isolates (CAU 1497, CAU 1499, 18N1A7, CAU 1496, isolation. The pure culture of bacterial isolates were pre- and 18H3T14) of the family Planococcaceae; 2 isolates served at -80°C as a suspension in liquid culture media (LPB0217 and LPB0242) of the family Staphylococca- containing 10-20% glycerol (w/v) and also preserved by ceae. In the order Lactobacillales, the 15 isolates further lyophilization. The 35 bacterial species designations, iso- grouped into 5 bacterial families; 1 isolate (LPB0228) lation sources, culture media, and incubation conditions of the family Aerococcaceae; 2 isolates (CAU 1515 and are summarized in Table 1. LPB0216) of the family Enterococcaceae; 4 isolates The 16S rRNA gene was amplified and sequenced using (CAU 1514, LPB0236, LPB0237, and LPB0238) of the standard PCR and Sanger sequencing methods with uni- family Lactobacillaceae; 3 isolates (LPB0215, LPB0235, versal primers 27F and 1492R. The determined sequences and LPB0239) of the family Leuconostocaceae; 5 iso- November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 339 2 days 3 days 2 days 3 days 2 days - - - - - condition Incubation 30℃, 2 days 30℃, 2 30℃, 2 days 30℃, 3 days 37℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 2 days 30℃, 3 days 30℃, 1 30℃, 2 30℃, 2 days 30℃, 3 days 30℃, 3 days 25℃, 3 days 25℃, 4 days 25℃, 3 days 30℃, 1 25℃, 4 days 30℃, 1 MH GY E MH AB TSY MA R2A MH R2A GY E MA MH R2A R2A L B R2A R2A MA R2A MA Medium ) ) ) ) ) ) ) ) ) ) ) ) ) ) ʺ ʺ ʺ ) ) ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ) ʹʺ ʺ 2.4 30.8 ʹ ʹ 44.2 58.2 59.21 ʺ ) 22.7 52.8 41.2 20.8 47.1 57.3 24.6 26 42.9 20.8 ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ 39.53 17.93 ʹ ʹ ; E 127°44.62 ; E 127°59 ; E 130°53 ; E 129°09 ; E 127°01 ; E 126°18 ; E 127°19 ; E 127°15 ; E 127°19 ; E 126°55 ; E 126°47 ; E 127°20 ; E 126°45 ; E 126°24 ; E 127°19 ʺ ʺ ʺ ; E 127°23 ; E 126°43 Soil Soil Soil Soil Soil Soil Soil Soil Soil Soil ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ Moss Wood (G P S) L ichen Sea salt Sea sand 59 Sea water ʹ Mouse gut (unknown) (unknown) 32.6 02.4 57.6 22.5 08.9 37.5 22.0 49.2 07.4 03.1 37.5 ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ 14.92 57.4 22.96 6.73 47.44 Mine drainage Mine drainage ʹ ʹ ʹ ʹ ʹ Human sputum Isolation source (N33°32 (N38°5 (N37°0 (N35°09 (N37°32 (N36°44 (N36°21 (N37°20 (N37°47 (N33°25 (N36°24 (N36°13 (N37°32 (N37°47 (N36°23 (N37°39 (N37°29 (N37°34 ʹ 42.78 ʺ ; E 126°59 (%)

98.7 99.0 99.7 99.2 99.8 99.9 99.9 98.9 99.2 99.7 99.2 99.2 99.6 98.8 99.9 99.6 99.6 99.4 99.1 99.2 16S rRNA 16S rRNA similarity gene sequence species. Most closely related species Firmicutes Paenibacillus methanolicus lasiicapitis Psychrobacillus psychrodurans Psychrobacillus Rummelibacillus pycnus Solibacillus isronensis Staphylococcus argenteus Staphylococcus lentus Anaerobacillus macyae Anaerobacillus Bacillus cucumis Bacillus psychrosaccharolyticus Paenibacillus anaericanus Paenibacillus lautus Paenibacillus polysaccharolyticus Paenibacillus salinicaeni Paenibacillus shirakamiensis Paenibacillus silage Paenibacillus tibetensis Bacillus solisilvae Halobacillus dabaenesis Lysinibacillus contaminans Lysinibacillus NIBR ID NIBRBAC000502511 NIBRBAC000502361 NIBRBAC000502551 NIBRBAC000502369 NIBRBAC000502547 NIBRBAC000502414 NIBRBAC000502440 NIBRBAC000502507 NIBRBAC000502569 NIBRBAC000502331 NIBRBAC000502535 NIBRBAC000502370 NIBRBAC000502542 NIBRBAC000502557 NIBRBAC000502516 NIBRBAC000502403 NIBRBAC000502333 NIBRBAC000502432 NIBRBAC000502365 NIBRBAC000502362 Strain designation HM F 7158 CA U 1499 18N1A7 CA U 1496 18H3T14 LP B0217 LP B0242 HM F 5940 GA079 18JY44-4 18S2 L 7 CA U 1503 18H7A6 MMS18-CY120 HM F 7562 GY1 18JY2-4 LP B0245 CA U 1495 CA U 1497 F amily

Strain information and taxonomic affiliation of the 35 unrecorded Staphylococcaceae Bacillaceae Paenibacillaceae Planococcaceae Bacillales Order Table 1. Table 340 Journal of Species Research Vol. 8, No. 4 2 days 2 days - - condition Incubation 37℃, 3 days 37℃, 3 days 37℃, 3 days 37℃, 3 days 37℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 3 days 30℃, 3 days 37℃, 3 days 37℃, 1 37℃, 1 TSY TSY TSY TSY TSY AB MRS MRS MRS MRS MRS MRS TSY MRS MRS Medium ) ) ) ) ) ) ) ) ) ) ) ) ) ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ) ) ʺ ʺ ʺ ʺ ʺ ʺ 59.21 59.21 59.21 59.21 59.21 40.82 40.82 40.82 59.21 32.6 32.6 ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ 39.53 39.53 39.53 39.53 ʹ ʹ ʹ ʹ ; E 126°59 ; E 126°59 ; E 126°59 ; E 126°59 ; E 126°59 ; E 127°27 ; E 127°27 ; E 127°27 ; E 126°59 ; E 126°57 ; E 126°57 ; E 127°23 ; E 127°23 ; E 127°23 ; E 127°23 ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ ʺ (G P S) Mouse gut Mouse gut Mouse gut Mouse gut Mouse gut Mouse gut Mouse gut 20.1 20.1 Human feces Human feces ʹ ʹ 57.4 57.4 57.4 57.4 ʹ ʹ ʹ ʹ Human sputum Human sputum Human sputum Human sputum Human sputum Human sputum 42.78 42.78 42.78 42.78 42.78 57.54 57.54 57.54 42.78 Isolation source ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ ʹ (N37°30 (N37°30 (N36°23 (N36°23 (N36°23 (N36°23 (N37°34 (N37°34 (N37°34 (N37°34 (N37°34 (N36°19 (N36°19 (N36°16 (N37°34 (%)

100 100 100 100 99.8 99.3 99.8 99.6 99.9 99.9 99.6 99.9 99.9 99.3 99.8 16S rRNA 16S rRNA similarity gene sequence whileyi . subsp. subsp species. Most closely related species Firmicutes Streptococcus cristatus Streptococcus gordonii Streptococcus mutans Streptococcus sanguinis Streptococcus Lactobacillus paracasei cibaria Weissella anginosus Streptococcus Enterococcus lactis Enterococcus Lactobacillus murinus Lactobacillus ruminis Pediococcus acidilactici confuse Weissella paramesenteroides Weissella Granulicatella adiacens avium Enterococcus tolerans NIBR ID NIBRBAC000502416 NIBRBAC000502418 NIBRBAC000502421 NIBRBAC000502422 NIBRBAC000502381 NIBRBAC000502437 NIBRBAC000502419 NIBRBAC000502413 NIBRBAC000502429 NIBRBAC000502427 NIBRBAC000502428 NIBRBAC000502426 NIBRBAC000502430 NIBRBAC000502439 NIBRBAC000502382 Strain designation LP B0219 LP B0221 LP B0225 LP B0227 CA U 1514 LP B0215 LP B0222 LP B0216 LP B0238 LP B0236 LP B0237 LP B0235 LP B0239 LP B0228 CA U 1515 F amily

Strain information and taxonomic affiliation of the 35 unrecorded Continued. Lactobacillaceae Leuconostocaceae Streptococcaceae Aerococcaceae Enterococcaceae Lactobacillales Order Table 1. 1. Table Table Anaerobe basal agar. AB, agar; L B, uria-Bertani Yeast Soy Trypticase TSY, E xtract agar; MRS, L actobacilli MRS Yeast Abbreviations: MA, Marine agar; MH, Mueller Hinton GY E , Glucose November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 341

Fig. 1. Neighbor-joining tree based on 16S rRNA gene sequences showing the phylogenetic relationships between the strains isolated in this study and their relatives of (A) the order Bacillales and (B) the order Lactobacillales. Filled circles indicate that the corresponding nodes (groupings) were also recovered in the maximum likelihood tree. Bootstrap values (>70%) are shown above nodes. Escherichia coli ATCC 11775T (X80725) was used as an outgroup. lates (LPB0219, LPB0221, LPB0222, LPB0225, and species). LPB0227) of the family Streptococcaceae. These unrecorded species showed diverse cellular mor- At the genus level, 9 genera were found in the order phologies including rods, short rods, cocci, or ovoid (Fig. Bacillales; Anaerobacillus (1 species), Bacillus (3 spe- 2). The colony colors were white, cream, yellow, orange, cies), Halobacillus (1 species), Paenibacillus (8 species), or red. Most species were positive at Gram-staining, but a Lysinibacillus (1 species), Psychrobacillus (2 species), minor portion was stained Gram-negative. All species re- Rummeliibacillus (1 species), Solibacillus (1 species), and ported here grew aerobically, but some species also grew Staphylococcus (2 species). Six genera were found in the at anaerobic condition indicating their facultative anaer- order Lactobacillales; Enterococcus (2 species), Granuli- obic respiration. The enzymatic and physiological prop- catella (1 species), Lactobacillus (3 species), Pediococcus erties examined by API kits were also variable depending (1 species), Streptococcus (5 species), and Weissella (3 on species. The features of oxidase, catalase, nitrate re- 342 Journal of Species Research Vol. 8, No. 4

Fig. 1. Continued. duction, indole production, glucose fermentation, arginine 35 species of 15 genera belonging to the phylum Firmic- dihydrolase, urease, β-galactosidase, gelatin hydrolysis, utes were found and examined taxonomically. The pres- esculin hydrolysis, and utilization or acidification of car- ence of the above mentioned Firmicutes species reported bon sources of each species are described in the proto- in overseas, but not in Korea. Thus, the following 35 spe- logues. cies are reported as unrecorded species in Korea and their From the natural and artificial environments in Korea, phenotypic properties are given. November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 343

A (1) (2) (3) (4) (5)

(6) (7) (8) (9) (10)

(11) (12) (13) (14) (15)

(16) (17) (18) (19) (20)

Fig. 2. Transmission electron micrographs or scanning electron micrographs of cells of (A) the order Bacillales and (B) the order Lactoba- cillales. Strains: 1, HMF5940; 2, GA079; 3, 18JY44-4; 4, LPB0245; 5, CAU 1495; 6, 18S2L7; 7, CAU 1503; 8, HMF7158; 9, 18H7A6; 10, MS18-CY120; 11, HMF7562; 12, GY1; 13, 18JY2-4; 14, CAU 1497; 15, CAU 1499; 16, 18N1A7; 17, CAU1496; 18, 18H3T14; 19, LPB0217; 20, LPB0242; 21, CAU 1515; 22, CAU 1514; 23, LPB0228; 24, LPB0216; 25, LPB0238; 26, LPB0236; 27, LPB0237; 28, LPB0215; 29, LPB0235; 30, LPB0239; 31, LPB0222; 32, LPB0219; 33, LPB0221; 34, LPB0225; 35, LPB0227.

Description of Anaerobacillus macyae HMF5940 MK123415. Cells are Gram-staining-positive, non-flagellated, rod- Description of Bacillus cucumis GA079 shaped, and aerobic. Colonies are circular, smooth, convex, and white colored after 3 days on MA at 30℃. Pos- Cells are Gram-staining-negative, non-flagellated, rod- itive for oxidase. Hydrolyses esculin, but not gelatin. Re- shaped, and aerobic. Colonies are circular, glistening, duces nitrate to nitrite. Cannot produce acid from glucose. convex, and cream colored after 3 days on R2A at 30℃. Does not produce indole. Possesses activity of β-galacto- Positive for oxidase. Hydrolyses esculin, but not gelatin. sidase, but not arginine dihydrolase or urease. Does not Reduces nitrate to nitrite. Cannot produce acid from glu- assimilate glucose, arabinose, mannose, mannitol, malt- cose. Does not produce indole. Possesses activity of β-ga- ose, N-acetyl-glucosamine, potassium gluconate, capric lactosidase, but not arginine dihydrolase or urease. Assim- acid, adipic acid, malic acid, trisodium citrate, or pheny- ilates glucose, arabinose, mannitol, maltose, N-acetyl-glu- lacetic acid. Strain HMF5940 (=NIBRBAC000502507) cosamine, potassium gluconate, but not mannose, capric was isolated from sea salt, Taean, Korea. The GenBank acid, adipic acid, malic acid, trisodium citrate, or phe- accession number for the 16S rRNA gene sequence is nylacetic acid. Strain GA079 (=NIBRBAC000502569) 344 Journal of Species Research Vol. 8, No. 4

B (21) (22) (23) (24) (25)

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

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

Fig. 2. Continued. was isolated from soil, Daejeon, Korea. The GenBank Negative for oxidase. Does not hydrolyze esculin and accession number for the 16S rRNA gene sequence is gelatin. Does not reduce nitrate to nitrite. Cannot produce MK135794. acid from glucose. Does not produce indole. Does not possess activity of β-galactosidase, arginine dihydrolase, Description of Bacillus psychrosaccharolyticus or urease. Weakly assimilates glucose, mannose, man- 18JY44-4 nitol, but not arabinose, maltose, N-acetyl-glucosamine, Cells are Gram-staining-positive, non-flagellated, coc- potassium gluconate, capric acid, adipic acid, malic acid, ci-shaped, and aerobic. Colonies are circular, convex, trisodium citrate, or phenylacetic acid. Strain LPB0245 smooth, and yellow colored after 4 days on R2A at 25℃. ( =NIBRBAC000502432) was isolated from soil, Ul- Hydrolyses esculin and gelatin. Reduces nitrate to nitrite. leung-do, Korea. The GenBank accession number for the Cannot produce acid from glucose. Does not produce 16S rRNA gene sequence is MH991823. indole. Possesses activity of β-galactosidase, but not arginine dihydrolase or urease. Assimilates glucose, arabi- Description of Enterococcus avium CAU 1515 nose, mannose, mannitol, maltose, N-acetyl-glucosamine, Cells are Gram-staining-positive, non-flagellated, potassium gluconate, adipic acid, but not capric acid, ovoid-shaped, and aerobic. Colonies are circular, convex, malic acid, trisodium citrate, or phenylacetic acid. Strain smooth, shiny, opaque, and white colored after 1-2 days 18JY44-4 ( =NIBRBAC000502331) was isolated from on MRS at 37℃. Hydrolyses esculin, but not gelatin. soil, Ulsan, Korea. The GenBank accession number for Does not reduce nitrate to nitrite. Can produce acid from the 16S rRNA gene sequence is MH934918. glucose. Does not produce indole. Does not possess activity of β-galactosidase, arginine dihydrolase, or urease. Description of Bacillus solisilvae LPB0245 Assimilates glucose, mannose, mannitol, N-acetyl-glucos- Cells are Gram-staining-positive, non-flagellated, rod- amine (weakly), maltose (weakly), and potassium gluco- shaped, and aerobic. Colonies are filamentous, rough, nate (weakly), but not arabinose, capric acid, adipic acid, convex, and white colored after 3 days on R2A at 25℃. malic acid, trisodium citrate, or phenylacetic acid. Strain November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 345

CAU 1515 (=NIBRBAC000502382) was isolated from Description of Lactobacillus murinus LPB0238 human feces, Seoul, Korea. The GenBank accession num- Cells are Gram-staining-positive, non-flagellated, rod- ber for the 16S rRNA gene sequence is MK182809. shaped, and facultative anaerobic. Colonies are irregular, undulate, wrinkled, flat, and creamy white colored after Description of Enterococcus lactis LPB0216 3 days on MRS at 30℃. Negative for oxidase. Produces Cells are Gram-staining-positive, non-flagellated, coc- indole. Does not possess activity of urease. Hydrolyses coid-shaped, and facultative anaerobic. Colonies are cir- esculin, but not gelatin. Weakly acidifies glucose, lac- cular, entire, smooth, convex, and white colored after 3 tose, saccharose, maltose, xylose, arabinose, cellobiose, days on AB at 30℃. Negative for oxidase. Does not pro- mannose, melezitose, raffinose, rhamnose, and trehalose, duce indole. Does not possess activity of urease. Hydro- but not mannitol, salicin, glycerol, or sorbitol. Strain lyses esculin, but not gelatin. Acidifies glucose, mannitol, LPB0238 ( =NIBRBAC000502429) was isolated from lactose, saccharose, maltose, salicin, arabinose, cellobi- mouse gut, Daejeon University, Korea. The GenBank ose, raffinose, rhamnose (weakly), and trehalose (weakly), accession number for the 16S rRNA gene sequence is but not xylose, glycerol, mannose, melezitose, or sorbitol. MK002979. Strain LPB0216 (=NIBRBAC000502413) was isolated from mouse gut, Korea Institute of Oriental Medicine, Description of Lactobacillus paracasei subsp. tolerans Korea. The GenBank accession number for the 16S rRNA CAU 1514 gene sequence is MH997849. Cells are Gram-staining-positive, non-flagellated, rod- Description of Granulicatella adiacens LPB0228 shaped, and aerobic. Colonies are circular, raised, smooth, shiny, opaque, and white colored after 1-2 days on MRS Cells are Gram-staining-positive, non-flagellated, cocci- at 37℃. Hydrolyses esculin, but not gelatin. Does not re- shaped, and aerobic. Colonies are circular, entire, smooth, duce nitrate to nitrite. Produces acid from glucose. Does convex, and cream colored after 3 days on TSY at 37℃. not produce indole. Possesses activity of β-galactosidase Negative for oxidase. Weakly hydrolyses esculin, but not (weakly), but not arginine dihydrolase or urease. Assim- gelatin. Does not reduce nitrate to nitrite. Cannot produce ilates glucose, arabinose, mannose, mannitol, maltose, acid from glucose. Does not produce indole. Does not N-acetyl-glucosamine (weakly), potassium gluconate, possess activity of β-galactosidase, arginine dihydrolase, capric acid, adipic acid, malic acid (weakly), trisodium or urease. Does not assimilate glucose, arabinose, man- citrate, and phenylacetic acid. Strain CAU 1514 (=NI- nose, mannitol, maltose, N-acetyl-glucosamine, potassi- BRBAC000502381) was isolated from human feces, um gluconate, capric acid, adipic acid, malic acid, trisodi- Seoul, Korea. The GenBank accession number for the um citrate, or phenylacetic acid. Strain LPB0228 (=NI- 16S rRNA gene sequence is MK182810. BRBAC000502439) was isolated from human sputum, Seoul National University Hospital, Korea. The GenBank Description of Lactobacillus ruminis LPB0236 accession number for the 16S rRNA gene sequence is MH930448. Cells are Gram-staining-positive, non-flagellated, rod- shaped, and facultative anaerobic. Colonies are circular, Description of Halobacillus dabanensis CAU 1495 entire, smooth, convex, and white colored after 3 days on MRS at 30℃. Negative for oxidase. Does not produce Cells are Gram-staining-positive, non-flagellated, indole. Does not possess activity of urease. Hydrolyses rod-shaped, and aerobic. Colonies are circular, convex, esculin, but not gelatin. Acidifies glucose, mannitol, lac- - smooth, shiny, opaque, and pale orange colored after 1 2 tose, saccharose, maltose, salicin, arabinose (weakly), days on MA at 30℃. Hydrolyses gelatin, but not esculin. cellobiose, mannose, and raffinose, but not xylose, glyc- Does not reduce nitrate to nitrite. Cannot produce acid erol, melezitose, sorbitol, rhamnose, or trehalose. Strain from glucose. Does not produce indole. Does not pos- LPB0236 ( =NIBRBAC000502427) was isolated from sess activity of β-galactosidase, arginine dihydrolase, or mouse gut, Korea Institute of Oriental Medicine, Korea. urease. Does not assimilate glucose, arabinose, mannose, The GenBank accession number for the 16S rRNA gene mannitol, maltose, N-acetyl-glucosamine, potassium sequence is MK002977. gluconate, capric acid, adipic acid, malic acid, trisodium citrate, or phenylacetic acid. Strain CAU 1495 ( =NI- Description of Lysinibacillus contaminans CAU 1497 BRBAC000502365) was isolated from human sputum, Seoul National University Hospital, Korea. The GenBank Cells are Gram-staining-positive, non-flagellated, accession number for the 16S rRNA gene sequence is rod-shaped, and aerobic. Colonies are circular, convex, MK053895. smooth, shiny, opaque, and cream colored after 1-2 days 346 Journal of Species Research Vol. 8, No. 4 on MA at 30℃. Does not hydrolyze esculin and gela- ase. Does not assimilate glucose, arabinose, mannose, tin. Does not reduce nitrate to nitrite. Cannot produce mannitol, maltose, N-acetyl-glucosamine, potassium acid from glucose. Produce indole. Possesses activi- gluconate, capric acid, adipic acid, malic acid, trisodium ty of arginine dihydrolase and urease (weakly), but not citrate, or phenylacetic acid. Strain HMF7158 ( =NI- β-galactosidase. Assimilates trisodium citrate, but not BRBAC000502511) was isolated from moss, Yongin, glucose, arabinose, mannose, mannitol, maltose, N-ace- Korea. The GenBank accession number for the 16S rRNA tyl-glucosamine, potassium gluconate, capric acid, adipic gene sequence is MK123430. acid, malic acid, or phenylacetic acid. Strain CAU 1497 (=NIBRBAC000502362) was isolated from mouse mine Description of Paenibacillus polysaccharolyticus drainage, Gapyeong-gun, Korea. The GenBank accession 18H7A6 number for the 16S rRNA gene sequence is MK053917. Cells are Gram-staining-positive, non-flagellated, rod- shaped, and facultative anaerobic. Colonies are circular, Description of Paenibacillus anaericanus 18S2L7 flat, smooth, undulate, and white colored after 2 days on Cells are Gram-staining-positive, flagellated, rod- MH at 30℃. Positive for oxidase. Hydrolyses esculin shaped, and facultative anaerobic. Colonies are circular, and gelatin. Reduces nitrate to nitrite. Cannot produce convex, smooth, and white colored after 2 days on MH at acid from glucose. Does not produce indole. Possesses 30℃. Negative for oxidase. Hydrolyses esculin, but not activity of β-galactosidase, but not arginine dihydrolase gelatin. Reduces nitrate to nitrite. Cannot produce acid or urease. Assimilates glucose, arabinose, mannitol, malt- from glucose. Does not produce indole. Possesses ac- ose, but not mannose, N-acetyl-glucosamine, potassium tivity of β-galactosidase, but not arginine dihydrolase or gluconate, capric acid, adipic acid, malic acid, trisodi- urease. Does not assimilate glucose, arabinose, mannose, um citrate, or phenylacetic acid. Strain 18H7A6 (=NI- mannitol, maltose, N-acetyl-glucosamine, potassium BRBAC000502542) was isolated from soil, Gimpo, Ko- gluconate, capric acid, adipic acid, malic acid, trisodi- rea. The GenBank accession number for the 16S rRNA um citrate, or phenylacetic acid. Strain 18S2L7 ( =NI- gene sequence is MK129286. BRBAC000502535) was isolated form soil, Chungju, Korea. The GenBank accession number for the 16S rRNA Description of Paenibacillus salinicaenis gene sequence is MK129280. MMS18-CY120 Cells are Gram-staining-variable, non-flagellated, rod- Description of Paenibacillus lautus CAU 1503 shaped, and aerobic. Colonies are circular, convex, smoo Cells are Gram-staining-positive, non-flagellated, th, and cream colored after 3 days on R2A at 30℃. Neg- rod-shaped, and aerobic. Colonies are circular, convex, ative for oxidase. Hydrolyses esculin and gelatin. Does smooth, shiny, opaque, and cream colored after 2-3 days not reduce nitrate to nitrite. Cannot produce acid from on MA at 30℃. Does not hydrolyze esculin and gelatin. glucose. Does not produce indole. Possesses activity of Does not reduce nitrate to nitrite. Cannot produce acid β-galactosidase and urease, but not arginine dihydrolase. from glucose. Does not produce indole. Possesses activ- Assimilates glucose, arabinose, mannose, mannitol, malt- ity of urease (weakly), but not arginine dihydrolase or ose, N-acetyl-glucosamine, potassium gluconate, malic β-galactosidase. Does not assimilate glucose, arabinose, acid, trisodium citrate and phenylacetic acid, but not mannose, mannitol, maltose, N-acetyl-glucosamine, po- capric or and adipic acid. Strain MMS18-CY120 (=NI- tassium gluconate, capric acid, adipic acid, malic acid, BRBAC000502557) was isolated from soil, Cheong- trisodium citrate, or phenylacetic acid. Strain CAU 1503 yand-gun, Korea. The GenBank accession number for the (=NIBRBAC000502370) was isolated from mouse sea 16S rRNA gene sequence is MK089843. water, Seopjikoji, Jeju-do, Korea. The GenBank accession number for the 16S rRNA gene sequence is MK053922. Description of Paenibacillus salinicaenis HMF7562 Cells are Gram-staining-positive, flagellated, rod- Description of Paenibacillus methanolicus HMF7158 shaped, and aerobic. Colonies are circular, convex, Cells are Gram-staining-positive, non-flagellated, smooth, and white colored after 3 days on R2A at 30℃. rod-shaped, and aerobic. Colonies are circular, convex, Positive for oxidase. Hydrolyses esculin, but not gela- smooth, and white colored after 3 days on R2A at 30℃. tin. Does not reduce nitrate to nitrite. Cannot produce Positive for oxidase. Does not hydrolyze esculin and gela- acid from glucose. Does not produce indole. Possesses tin. Does not reduce nitrate to nitrite. Cannot produce acid activity of β-galactosidase, but not arginine dihydrolase from glucose. Does not produce indole. Dose not possess or urease. Assimilates glucose, maltose, N-acetyl-glucos- activity of arginine dihydrolase, β-galactosidase, or ure- amine, but not arabinose, mannose, mannitol, potassium November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 347 gluconate, capric acid, adipic acid, malic acid, trisodium Description of Psychrobacillus lasiicapitis CAU 1499 citrate, or phenylacetic acid. Strain HMF7562 ( =NI- Cells are Gram-staining-positive, non-flagellated, rod- BRBAC000502516) was isolated from wood, Daejeon, shaped, and aerobic. Colonies are circular, convex, smoo Korea. The GenBank accession number for the 16S rRNA th, shiny, opaque, and yellow colored after 2-3 days on gene sequence is MK123434. GYE at 30℃. Hydrolyses esculin, but not gelatin. Does not reduce nitrate to nitrite. Cannot produce acid from Description of Paenibacillus silageis GY1 glucose. Does not produce indole. Does not possess ac- Cells are Gram-staining-negative, non-flagellated, rod- tivity of β-galactosidase, arginine dihydrolase, or urease. shaped, and aerobic. Colonies are circular, convex, smoo Assimilates glucose, arabinose (weakly), mannitol (weak- th, and white colored after 3 days on LB at 25℃. Positive ly), maltose (weakly), potassium gluconate, capric acid for oxidase. Hydrolyses esculin, but not gelatin. Reduces (weakly), adipic acid (weakly), and trisodium citrate, but nitrate to nitrite. Cannot produce acid from glucose. Does not mannose, N-acetyl-glucosamine, malic acid, or pheny- not produce indole. Possesses activity of β-galactosidase, lacetic acid. Strain CAU 1499 (=NIBRBAC000502361) but not arginine dihydrolase or urease. Assimilates arab- was isolated from sea sand, Busan, Korea. The GenBank inose, mannose, mannitol and maltose, but not glucose, accession number for the 16S rRNA gene sequence is N-acetyl-glucosamine, potassium gluconate, capric acid, MK053919. adipic acid, malic acid, trisodium citrate, or phenylacetic acid. Strain GY1 (=NIBRBAC000502403) was isolated Description of Psychrobacillus psychrodurans 18N1A7 from lichen, Gimnyeong, Jeju-do, Korea. The GenBank Cells are Gram-staining-positive, flagellated, rod- accession number for the 16S rRNA gene sequence is shaped, and aerobic. Colonies are circular, convex, smoo MK201775. th, and white colored after 2 days on MH at 30℃. Neg- ative for oxidase. Hydrolyses esculin, but not gelatin. Description of Paenibacillus tibetensis 18JY2-4 Does not reduce nitrate to nitrite. Cannot produce acid Cells are Gram-staining-negative, non-flagellated, rod- from glucose. Does not produce indole. Possesses ac- shaped, and aerobic. Colonies are circular, convex, smoo tivity of β-galactosidase, but not arginine dihydrolase or th, and yellow colored after 4 days on R2A at 25℃. Hy- urease. Does not assimilate glucose, arabinose, mannose, drolyses esculin, but not gelatin. Does not reduce nitrate mannitol, maltose, N-acetyl-glucosamine, potassium to nitrite. Cannot produce acid from glucose. Does not gluconate, capric acid, adipic acid, malic acid, trisodi- produce indole. Does not possess activity of β-galactosi- um citrate, or phenylacetic acid. Strain 18N1A7 (=NI- dase, arginine dihydrolase, or urease. Assimilates glucose, BRBAC000502551) was isolated from around the river N-acetyl-glucosamine, and maltose, but not arabinose, soil, Hwachon-gun, Korea. The GenBank accession num- mannose, mannitol, potassium gluconate, capric acid, ber for the 16S rRNA gene sequence is MK129297. adipic acid, malic acid, trisodium citrate, or phenylacetic acid. Strain 18JY2-4 ( =NIBRBAC000502333) was Description of Rummeliibacillus pycnus CAU 1496 isolated from soil, Ulleung-gun, Korea. The GenBank Cells are Gram-staining-positive, non-flagellated, oval accession number for the 16S rRNA gene sequence is rod-shaped, and aerobic. Colonies are circular, flat, smoo MH934921. th, lobate margin, shiny, and cream colored after 1-2 days on GYE at 30℃. Does not hydrolyze esculin or gelatin. Description of Pediococcus acidilactici LPB0237 Does not reduce nitrate to nitrite. Cannot produce acid from glucose. Produce indole. Dose not possess activity of Cells are Gram-staining-positive, non-flagellated, coc- β-galactosidase, arginine dihydrolase, or urease. Does not coid-shaped, and facultative anaerobic. Colonies are circu- assimilate glucose, arabinose, mannose, mannitol, malt- lar, entire, smooth, convex, and creamy white colored after ose, N-acetyl-glucosamine, potassium gluconate, capric 3 days on MRS at 30℃. Negative for oxidase. Does not acid, adipic acid, malic acid, trisodium citrate, or pheny- produce indole. Does not possess activity of urease. Hy- lacetic acid. Strain CAU 1496 (=NIBRBAC000502369) drolyses esculin, but not gelatin. Acidifies glucose, man- was isolated from mine drainage, Gapyeong-gun, Korea. nitol, lactose, saccharose, maltose, salicin, xylose, arab- The GenBank accession number for the 16S rRNA gene inose, glycerol (weakly), cellobiose, mannose, raffinose, sequence is MK053894. sorbitol (weakly), rhamnose, and trehalose, but not melezitose. Strain LPB0237 ( =NIBRBAC000502428) Description of Solibacillus isronensis 18H3T14 was isolated from mouse gut, Korea Institute of Oriental Medicine, Korea. The GenBank accession number for the Cells are Gram-staining-positive, non-flagellated, rod- 16S rRNA gene sequence is MK002978. shaped, and aerobic. Colonies are circular, raised, smooth, 348 Journal of Species Research Vol. 8, No. 4 and white colored after 2 days on MH at 30℃. Negative produce acid from glucose. Does not produce indole. Pos- for oxidase. Does not hydrolyze esculin or gelatin. Does sesses activity of β-galactosidase (weakly) and arginine not reduce nitrate to nitrite. Cannot produce acid from dihydrolase, but not urease. Does not assimilate glucose, glucose. Does not produce indole. Dose not possess ac- arabinose, mannose, mannitol, maltose, N-acetyl-glu- tivity of β-galactosidase, arginine dihydrolase, or ure- cosamine, potassium gluconate, capric acid, adipic acid, ase. Assimilates potassium gluconate, but not glucose, malic acid, trisodium citrate, or phenylacetic acid. Strain arabinose, mannose, mannitol, maltose, N-acetyl-glu- LPB0222 ( =NIBRBAC000502419) was isolated from cosamine, capric acid, adipic acid, malic acid, trisodium human sputum, Seoul National University Hospital, Ko- citrate, or phenylacetic acid. Strain 18H3T14 ( =NI- rea. The GenBank accession number for the 16S rRNA BRBAC000502547) was isolated from around the river gene sequence is MH930443. soil, Seoul, Korea. The GenBank accession number for the 16S rRNA gene sequence is MK129290. Description of Streptococcus cristatus LPB0219 Cells are Gram-staining-positive, non-flagellated, cocci- Description of Staphylococcus argenteus LPB0217 shaped, and facultative anaerobic. Colonies are circu- Cells are Gram-staining-positive, non-flagellated, coc- lar, entire, smooth, convex, and cream colored after 3 coid-shaped, and facultative anaerobic. Colonies are cir- days on TSY at 37℃. Negative for oxidase. Hydrolyses cular, entire, smooth, convex, and creamy white colored esculin (weakly), but not gelatin. Does not reduce nitrate after 3 days on ABat 30℃. Negative for oxidase. Does to nitrite. Cannot produce acid from glucose. Does not not produce indole. Possesses activity of urease. Does produce indole. Does not possess activity of β-galactosi- not hydrolyze esculin or gelatin. Acidifies glucose, lac- dase, arginine dihydrolase, or urease. Does not assimilate tose, saccharose, maltose, mannose, and trehalose, but glucose, arabinose, mannose, mannitol, maltose, N-ace- not mannitol, salicin, xylose, arabinose, glycerol, cello- tyl-glucosamine, potassium gluconate, capric acid, adipic biose, melezitose, raffinose, sorbitol, or rhamnose. Strain acid, malic acid, trisodium citrate, or phenylacetic acid. LPB0217 ( =NIBRBAC000502414) was isolated from Strain LPB0219 (=NIBRBAC000502416) was isolated mouse gut, Korea Institute of Oriental Medicine, Korea. from human sputum, Seoul National University Hospital, The GenBank accession number for the 16S rRNA gene Korea. The GenBank accession number for the 16S rRNA sequence is MH997860. gene sequence is MH930437.

Description of Staphylococcus lentus LPB0242 Description of Streptococcus gordonii LPB0221 Cells are Gram-staining-positive, non-flagellated, cocci- Cells are Gram-staining-positive, non-flagellated, cocci- shaped, and facultative anaerobic. Colonies are circular, shaped, and facultative anaerobic. Colonies are circular, entire, smooth, convex, and cream colored after 3 days on entire, smooth, convex, and cream colored after 3 days on TSY at 37℃. Positive for oxidase. Hydrolyses esculin, TSY at 37℃. Negative for oxidase. Hydrolyses esculin but not gelatin. Reduces nitrate to nitrite. Cannot produce (weakly), but not gelatin. Does not reduce nitrate to ni- acid from glucose. Does not produce indole. Possesses trite. Can produce acid from glucose. Does not produce activity of β-galactosidase, but not arginine dihydrolase indole. Does not possess activity of β-galactosidase, argi- or urease. Does not assimilate glucose, arabinose, man- nine dihydrolase, or urease. Does not assimilate glucose, nose, mannitol, maltose, N-acetyl-glucosamine, potassi- arabinose, mannose, mannitol, maltose, N-acetyl-glu- um gluconate, capric acid, adipic acid, malic acid, trisodi- cosamine, potassium gluconate, capric acid, adipic acid, um citrate, or phenylacetic acid. Strain LPB0242 (=NI- malic acid, trisodium citrate, or phenylacetic acid. Strain BRBAC000502440) was isolated from human sputum, LPB0221 ( =NIBRBAC000502418) was isolated from Seoul National University Hospital, Korea. The GenBank human sputum, Seoul National University Hospital, Ko- accession number for the 16S rRNA gene sequence is rea. The GenBank accession number for the 16S rRNA MK128421. gene sequence is MH930442.

Description of Streptococcus anginosus subsp. whileyi Description of Streptococcus mutans LPB0225 LPB0222 Cells are Gram-staining-positive, non-flagellated, cocci- Cells are Gram-staining-positive, non-flagellated, cocci- shaped, and facultative anaerobic. Colonies are circular, shaped, and facultative anaerobic. Colonies are circular, entire, smooth, convex, and cream colored after 3 days on entire, smooth, convex, and cream colored after 3 days on TSY at 37℃. Negative for oxidase. Hydrolyses esculin, TSY at 37℃. Negative for oxidase. Hydrolyses esculin but not gelatin. Does not reduce nitrate to nitrite. Can- and gelatin. Does not reduce nitrate to nitrite. Cannot not produce acid from glucose. Does not produce indole. November 2019 Baek et al. Unreported Firmicutes bacteria in Korea 349

Does not possess activity of β-galactosidase, arginine LPB0235 ( =NIBRBAC000502426) was isolated from dihydrolase, or urease. Does not assimilate glucose, arabi- mouse gut, Korea Institute of Oriental Medicine, Korea. nose, mannose, mannitol, maltose, N-acetyl-glucosamine, The GenBank accession number for the 16S rRNA gene potassium gluconate, capric acid, adipic acid, malic acid, sequence is MK002976. trisodium citrate, or phenylacetic acid. Strain LPB0225 ( =NIBRBAC000502421) was isolated from human Description of Weissella paramesenteroides LPB0239 sputum, Seoul National University Hospital, Korea. The Cells are Gram-staining-positive, non-flagellated, coc- GenBank accession number for the 16S rRNA gene se- coid-shaped, and facultative anaerobic. Colonies are quence is MF930444. circular, entire, smooth, convex, and creamy white colored after 3 days on MRS at 30℃. Negative for oxidase. Description of Streptococcus sanguinis LPB0227 Does not produce indole. Dose not possess activity of Cells are Gram-staining-positive, non-flagellated, cocci- urease. Does not hydrolyze esculin and gelatin. Acidifies shaped, and facultative anaerobic. Colonies are circular, glucose, lactose (weakly), saccharose, maltose, salicin entire, smooth, convex, and cream colored after 3 days on (weakly), xylose (weakly), arabinose, raffinose and tre- TSY at 37℃. Negative for oxidase. Hydrolyses esculin halose, but not mannitol, glycerol, cellobiose, mannose, (weakly), but not gelatin. Does not reduce nitrate to ni- melezitose, sorbitol, or rhamnose. Strain LPB0239 (=NI- trite. Can produce acid from glucose. Does not produce BRBAC000502430) was isolated from mouse gut, Dae- indole. Does not possess activity of β-galactosidase, argi- jeon University, Korea. The GenBank accession number nine dihydrolase, or urease. Does not assimilate glucose, for the 16S rRNA gene sequence is MK002980. arabinose, mannose, mannitol, maltose, N-acetyl-glucosamine, potassium gluconate, capric acid, adipic acid, malic acid, trisodium citrate, or phenylacetic acid. Strain Acknowledgements LPB0227 ( =NIBRBAC000502422) was isolated from human sputum, Seoul National University Hospital, Ko- This study was supported by the research grant “The rea. The GenBank accession number for the 16S rRNA Survey of Korean Indigenous Species” from the National gene sequence is MH930446. Institute of Biological Resources of the Ministry of Envi- ronment in Korea. Description of Weissella cibaria LPB0215 Cells are Gram-staining-positive, non-flagellated, short rod-shaped, and facultative anaerobic. Colonies are circu- References lar, entire, smooth, convex, and white colored after 3 days on MRS at 30℃. Negative for oxidase. Does not produce Bäckhed, F., H. Ding, T. Wang, L.V. Hooper, G.Y. Koh, A. indole. Does not possess activity of urease. Hydroly- Nagy and J.I. Gordon. 2004. The gut microbiota as an en- ses esculin, but not gelatin. Acidifies glucose, mannitol, vironmental factor that regulates fat storage. Proceedings lactose, saccharose, maltose, salicin, xylose, arabinose, of the National Academy of Sciences. USA 10144:15718- cellobiose, mannose, raffinose, rhamnose, and trehalose, 15723. but not glycerol, melezitose, or sorbitol. Strain LPB0215 Gibbons, N. and R. Murray. 1978. Proposals concerning the (=NIBRBAC000502437) was isolated from mouse gut, higher taxa of bacteria. International Journal of Systematic Korea Institute of Oriental Medicine, Korea. The Gen- and Evolutionary Microbiology 28:1-6. Bank accession number for the 16S rRNA gene sequence Jeffery, I.B., P.W. O’toole, L. Öhman, M.J. Claesson, J. is MH997848. Deane, E.M. Quigley and M. Simrén. 2012. An irritable bowel syndrome subtype defined by species-specific alter- Description of Weissella confusa LPB0235 ations in faecal microbiota. Gut 61:997-1006. Jeon, Y.-S., K. Lee, S.-C. Park, B.-S. Kim, Y.-J. Cho, S.-M. Cells are Gram-staining-positive, non-flagellated, short Ha and J. Chun. 2014. EzEditor: a versatile sequence rod-shaped, and facultative anaerobic. Colonies are circu- alignment editor for both rRNA- and protein-coding lar, entire, smooth, convex, and white colored after 3 days genes. International Journal of Systematic and Evolution- on MRS at 30℃. Negative for oxidase. Produces indole. ary Microbiology 64:689-691. Does not possess activity of urease. Hydrolyses esculin, Kumar, S., G. Stecher and K. Tamura. 2016. MEGA7: molec- but not gelatin. Acidifies glucose, saccharose, maltose, ular evolutionary genetics analysis version 7.0 for bigger salicin, xylose, cellobiose, mannose, raffinose (weakly), datasets. Molecular Biology and Evolution 33:1870-1874. and trehalose (weakly), but not mannitol, lactose, arabi- Ley, R.E., F. Bäckhed, P. Turnbaugh, C.A. Lozupone, R.D. nose, glycerol, melezitose, sorbitol, or rhamnose. Strain Knight and J.I. Gordon. 2005. Obesity alters gut microbial 350 Journal of Species Research Vol. 8, No. 4

ecology. Proceedings of the National Academy of Scienc- systematic bacteriology: Volume 3: The Firmicutes (Vol. es. USA 102:11070-11075. 3): Springer Science & Business Media. Rajilić-Stojanović, M., E. Biagi, H.G. Heilig, K. Kajander, Yoon, S.-H., S.-M. Ha, S. Kwon, J. Lim, Y. Kim, H. Seo R.A. Kekkonen, S. Tims and W.M. de Vos. 2011. Global and J. Chun. 2017. Introducing EzBioCloud: a taxonom- and deep molecular analysis of microbiota signatures in ically united database of 16S rRNA gene sequences and fecal samples from patients with irritable bowel syndrome. whole-genome assemblies. International Journal of Sys- Gastroenterology 141:1792-1801. tematic and Evolutionary Microbiology 67:1613-1617. Turnbaugh, P.J., M. Hamady, T. Yatsunenko, B.L. Cantarel, A. Duncan, R.E. Ley and J.P. Affourtit. 2009. A core gut microbiome in obese and lean twins. Nature 457:480-484. Submitted: September 20, 2019 De Vos, P., G. Garrity, D. Jones, N.R. Krieg, W. Ludwig, F.A. Revised: October 17, 2019 Rainey and W.B. Whitman. 2011. Bergey’s manual of Accepted: October 18, 2019