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An actinobacterial strain designated MMS20-HV4-12 T , displaying a high hydrolytic potential for various substrates, was isolated from a riverside soil sample and characterized by polyphasic taxonomic analysis. Growth occurred at 10-37 °C (optimum, 30°C), with NaCl concentrations of 0-4 % (optimum, 0 %) and at pH 7-9 (optimum, pH 8). MMS20-HV4-12 T was catalase-positive, oxidase-negative, rod-shaped and formed creamy white-coloured colonies. Based on the results of 16S rRNA gene sequence analysis, MMS20-HV4-12 T was found to be mostly related to the type strains of Nocardioides alpinus (98.3 % sequence similarity), Nocardioides furvisabuli (98.1 %) and Nocardioides zeicaulis (98.0 %). MMS20-HV4-12 T showed optimal growth on Reaoner's 2A agar, forming white-coloured colonies. The diagnostic polar lipid profile consisted of diphosphatidylglycerol, phosphatidylglycerol and phosphatidylinositol, the major fatty acids were iso-C 16 : 0 , C 17 : 1  ω8 c and 10-methyl-C 17 : 0 , the major isoprenoid quinone was MK-8(H 4 ), the diagnostic cell-wall sugar was galactose, and the cell-wall diamino acid was ll-diaminopimelic acid. The genome of MMS20-HV4-12 T was 4.47 Mbp in size with a G+C content of 72.9 mol%. The genome based analysis indicated low relatedness between MMS20-HV4-12 T and all compared species of Nocardioides , as the highest digital DNA-DNA hybridization and the orthologous average nucleotide identity values were 26.8 and 83.8% respectively. Based on genotypic, phenotypic and phylogenomic characterization, MMS20-HV4-12 T evidently represents a novel species of genus Nocardioides , for which the name Nocardioides okcheonensis sp. nov. (type strain=MMS20-HV4-12 T =KCTC 49651 T =LMG 32360 T ) is proposed.

A Gram-stain positive, aerobic, irregularly rod-shaped, non-spore-forming bacterium, designated as BN130099 T , was isolated from farmland soil sampled in Goesan-gun, Chungbuk, Republic of Korea. Phylogenetic analysis of its 16S rRNA gene sequence showed that the strain is closely related to Nocardioides pelophilus KACC 19192 T with 98.11 % similarity. The DNA G+C content of strain BN130099 T was 68.84 mol% (draft genome sequence). The genome sequence of BN130099 T displayed key enzymes involved in bioremediation of organic pollutants and biosynthetic clusters of saquayamycin. The strain contained ll-2,6-diaminopimelic acid in the cell-wall peptidoglycan and MK-8(H 4 ) as the major respiratory quinone. The predominant fatty acid was iso-C 16 : 0 . The major polar lipids were diphosphatidylglycerol, phosphatidylglycerol, phosphatidylcholine and phosphatidylinositol. The results of physiological and biochemical characterization allowed the phenotypic differentiation of strain BN130099 T from N. pelophilus KACC 19192 T . The strain represents a novel species of the genus Nocardioides , for which we propose the name Nocardioides humilatus sp. nov. The type strain is BN130099 T (=KCTC 49079 T =CCTCC AB 2018135 T ).

Two bacterial strains, designated MJB4 T and SJ7 T , were isolated from water samples collected from Jeongbang Falls on Jeju Island, Republic of Korea. Phylogenetic analysis of 16S rRNA gene sequences indicated that the two strains belonged to the genera Nocardioides and Hyunsoonleella , owing to their high similarities to Nocardioides jensenii DSM 29641 T (97.5 %) and Hyunsoonleella rubra FA042 T (96.3 %), respectively. These values are much lower than the gold standard for bacterial species (98.7 %). The average nucleotide identity values between strains MJB4 T , SJ7 T and the reference strains, Nocardioides jensenii DSM 29641 T , Nocardioides daejeonensis MJ31 T and Hyunsoonleella flava T58 T were 77.2, 75.9 and 75.4 %, respectively. Strains MJB4 T and SJ7 T and the type strains of the species involved in system incidence have average nucleotide identity and average amino acid threshold values of 60.1-82.6 % for the species boundary (95-96 %), which confirms that strains MJB4 T and SJ7 T represent two new species of genus Nocardioides and Hyunsoonleella , respectively. Based on phylogenetic and phenotypic data, strains MJB4 T and SJ7 T are considered to represent novel species of the genus Nocardioides and Hyunsoonleella , respectively, for which the names Nocardioides donggukensis sp. nov. (type strain MJB4 T =KACC 21724 T =NBRC 114402 T ) and Hyunsoonleella aquatilis sp. nov., (type strain SJ7 T =KACC 21715 T =NBRC 114486 T ) have been proposed.

In the present study, four bacterial strains, two (S-713 T and 406) isolated from faecal samples of Tibetan antelopes and the other two (S-531 T and 1598) from leaves of dandelion collected on the Qinghai-Tibet Plateau of PR China, were analysed using a polyphasic approach. All four isolates were aerobic, rod-shaped, non-motile, oxidase-negative, Gram-stain-positive and catalase-positive. According to four phylogenetic trees, strain pairs S-713 T /406 and S-531 T /1598 form two independent branches belonging to the genus Nocardioides , and are closest to Nocardioides lianchengensis , Nocardioides dokdonensis , Nocardioides salarius , Nocardioides marinisabuli , Nocardioides psychrotolerans and Nocardioides szechwanensis . Although sharing MK8-(H 4 ) as their major isoprenoid quinone, strains S-713 T and S-531 T contained C 18 : 1  ω 9 c (24.64 and 16.34 %) and iso-C 16 : 0 (9.74 and 29.38 %), respectively, as their main fatty acids, with remarkable differences in their biochemical profiles but only slight ones in their optimal growth conditions. The chromosomes of strains S-713 T and S-531 T were 4 207 844 bp (G+C content, 73.0 mol%) and 4 809 817 bp (G+C content, 72.5 mol%), respectively. Collectively, the two strain pairs represent two separate novel species of the genus Nocardioides , for which the names Nocardioides dongkuii sp. nov. and Nocardioides lijunqiniae sp. nov. are proposed, with S-713 T (=JCM 33698 T =CGMCC 4.7660 T ) and S-531 T (=JCM 33468 T =CGMCC 4.7659 T ) as the respective type strains.

A Gram-stain-positive, rod-shaped, strictly aerobic bacterial strain (Y6 T ) was isolated from a sewage sludge sample collected from a fisheries processing factory in Zhoushan, Zhejiang Province, PR China. The growth range of NaCl concentration was 0-6.0 % (w/v), with an optimum at 3.0 % (w/v). The temperature range for growth was 10-42 °C, with an optimum at 37 °C. The pH range for growth was pH 7.0-10.0, with an optimum at pH 9.0. Phylogenetic analysis based on 16S rRNA gene sequences revealed that strain Y6 T belonged to the genus Nocardioides and showed the highest sequence similarity of 97.8 % to Nocardioides jishulii dk3136 T . The average nucleotide identity and in silico DNA-DNA hybridization values between strain Y6 T and the reference strains were 76.9-81.2 % and 20.6-23.6 %, respectively. Chemotaxonomic analysis indicated that the sole respiratory quinone was MK-8(H 4 ) and the predominant cellular fatty acids were iso-C 16 : 0 , 10-methyl-C 17 : 0 and C 18 : 1  ω 9 c . The polar lipid profile was composed of diphosphatidylglycerol, phosphatidylglycerol, phosphatidylethanolamine, phosphatidylcholine, four unidentified phospholipids, three unidentified aminolipids and five unidentified lipids. The peptidoglycan was ll-2,6-diaminopimelic acid. On the basis of the phenotypic, genotypic, phylogenetic and chemotaxonomic features, strain Y6 T is considered to represent a novel species, for which the name Nocardioides malaquae sp. nov. is proposed. The type strain is Y6 T (=KCTC 49504 T =MCCC 1K04765 T ).

A polyphasic taxonomic approach was used to characterize three novel bacterial strains, designated as HDW12A T , HDW-15B T , and HDW15C T , isolated from the intestine of fish species Odontobutis interrupta or Siniperca scherzeri. All isolates were obligate aerobic, non-motile bacteria, and grew optimally at 30°C. Phylogenetic analysis based on 16S rRNA sequences revealed that strain HDW12A T was a member of the genus Nocardioides, and closely related to Nocardioides allogilvus CFH 30205 T (98.9% sequence identities). Furthermore, strains HDW15B T and HDW15C T were members of the genus Sphingomonas, and closely related to Sphingomonas lutea JS5 T and Sphingomonas sediminicola Dae 20 T (97.1% and 97.9% sequence identities), respectively. Strain HDW12A T contained MK-8 (H 4 ), and strains HDW15B T and HDW15C T contained Q-10 as the respiratory quinone. Major polar lipid components of strain HDW12A T were diphosphatidylglycerol, phosphatidylglycerol, and phosphatidylinositol, and those of strains HDW15B T and HDW15C T were sphingoglycolipid, diphosphatidylglycerol, phosphatidylglycerol, phosphatidylethanolamine, and phosphatidylcholine. The G + C content of strains HDW12A T , HDW15B T , and HDW15C T were 69.7, 63.3, and 65.5%, respectively. The results of phylogenetic, phenotypic, chemotaxonomic, and genotypic analyses suggest that strain HDW12A T represents a novel species within the genus Nocardioides, and strains HDW15B T and HDW15C T represent two novel species within the genus Sphingomonas. We propose the names Nocardioides piscis for strain HDW12A T (= KACC 21336 T = KCTC 49321 T = JCM 33670 T ), Sphingomonas piscis for strain HDW15B T (= KACC 21341 T = KCTC 72588 T = JCM 33738 T ), and Sphingomonas sinipercae for strain HDW15C T (= KACC 21342 T = KCTC 72589 T = JCM 33739 T ).

An aerobic, rod-shaped, Gram-stain-positive, actinobacterial strain, designated 1.0914 T , was isolated from a stalactite sample collected from a cave located in Guizhou Province, southwest PR China. Based on 16S rRNA gene sequence analysis, strain 1.0914 T shared highest similarities values with Nocardioides pelophilus CGMCC 4.7388 T (97.7 %), Nocardioides immobilis CCTCC AB 2017083 T (97.5 %) and Nocardioides silvaticus CCTCC AB 2018079 T (97.3 %) and values lower than 97.0 % to other members of the genus Nocardioides . Phylogenetic trees based on 16S rRNA gene sequences indicated that strain 1.0914 T formed an isolated branch with N. pelophilus CGMCC 4.7388 T , N. immobilis CCTCC AB 2017083 T and N. silvaticus CCTCC AB 2018079 T . The polar lipids contained phosphatidylglycerol, diphosphatidylglycerol, phosphatidylinositol and one unidentified phospholipid in the cellular membrane. The major fatty acids were identified as iso-C 16 : 0 , C 18 : 1  ω9 c , C 17 : 1  ω8 c and C 16 : 0 . The predominant respiratory quinone was MK-8(H 4 ) and ll-diaminopimelic acid was the diagnostic diamino acid in the cell-wall peptidoglycan. The genomic DNA G+C content was 71.1 mol%. The orthologous average nucleotide identiy values between N. pelophilus CGMCC 4.7388 T , N. immobilis CCTCC AB 2017083 T , N. silvaticus CCTCC 2018079 T and strain 1.0914 T were 82.3, 81.7 and 81.9 % respectively. DNA-DNA hybridization values between N. pelophilus CGMCC 4.7388 T , N. immobilis CCTCC AB 2017083 T , N. silvaticus CCTCC 2018079 T and strain 1.0914 T were 25.2, 24.6 and 24.5 % respectively. The phylogenetic, phenotypic and chemotaxonomic data supported the classification of strain 1.0914 T as representing a new species of Nocardioides , for which the name Nocardioides stalactiti sp. nov. is proposed. The type strain is 1.0914 T (=CCTCC AB 2018266 T =KCTC 49243 T ).

A novel actinobacterial strain, SB3-45 T , was isolated from soil of Cynanchum wilfordii rhizosphere, Jaecheon-si, Chungcheongbuk-do, Republic of Korea. Strain SB3-45 T , was Gram-stain-positive, aerobic and coccoid to short rod-shaped bacterium. Growth occurred at 4-37 °C (optimum 28 °C), pH 5-8 (optimum pH 7) and 0-2.5 % NaCl (optimum 0%). Phylogenetic analysis based on 16S rRNA gene sequence showed that strain SB3-45 T belonged to the genus Nocardioides and was closely related to Nocardioides opuntiae OS-21 T (96.2%) and Nocardioides panacihumi Gsoil 616 T (95.9%). ll-DAP as the diamino acid in the peptidoglycan and the menaquinone MK-8(H 4 ) as the predominant isoprenoid quinone were detected. The polar lipids of strain SB3-45 T were diphosphatidylglycerol, phosphatidylglycerol, phosphatidylinositol and unidentified phospholipid. The major cellular fatty acids (>5%) of strain SB3-45 T were iso-C 16 : 0 , C 18 : 1  ω9c and C 17 : 0 . Based on phylogenetic, physiological and chemotaxonomic characteristics, strain SB3-45 T represents a novel species of the genus Nocardioides, for which the name Nocardioides cynanchi sp.nov. is proposed. The type strain is SB3-45 T (=KCTC 49133 T =NBRC 114107 T ).

Two bacterial strains designated as W3-2-3 T and HKS04 T were isolated from mineral water and a soil sample, respectively, in the Republic of Korea. The 16S rRNA genes of the two strains shared a sequence similarity of 93.5 %. Phylogenetic analysis based on 16S rRNA gene sequences showed that strains W3-2-3 T and HKS04 T formed a distinct lineage within the genus Nocardioides of the family Nocardioidaceae (order Propionibacteriales ). The closely related species of strain W3-2-3 T were Nocardioides albidus (98.9 %), Nocardioides caeni (98.8 %), Nocardioides kongjuensis (98.6 %), Nocardioides aromaticivorans (98.5 %), Nocardioides nitrophenolicus (98.4 %), Nocardioides flava (98.2 %) and Nocardioides ginsengisoli (98.1 %). The closest species of strain HKS04 T was Nocardioides halotolerans (98.7 %). The genome sizes of strains W3-2-3 T and HKS04 T were 4741198 and 5 120341 bp, respectively. The genomic DNA G+C contents of strains W3-2-3 T and HKS04 T were 73.3 and 72.1 mol%, respectively. The main fatty acids of strain W3-2-3 T were C 17:1 ω 6 c and iso-C 16:0 and those of strain HKS04 T were iso-C 16:0 and iso-C 16:0 H. The main polar lipids of both strains were diphosphatidylglycerol and phosphatidylglycerol and the predominant respiratory quinone was MK-8(H 4 ), supporting the affiliation of these strains with the genus Nocardioides . Based on the results of biochemical, chemotaxonomic and phylogenetic analyses, two novel species, Nocardioides convexus W3-2-3 T (KACC 21211 T =LMG 31251 T ) and Nocardioides anomalus HKS04 T (KACC 18879 T =LMG 31249 T ), are proposed.

A Gram-stain-positive, non-motile, aerobic, rod-shaped, golden yellow-pigmented actinobacterium, designated strain YIM 123512 T , was isolated from soil sampled at Gaoligong Mountain, Yunnan Province, PR China. The strain grew at 10-40 °C (optimum, 28 °C), with 0-3.0 % (w/v) NaCl (optimum, 0-1.0 %) and at pH 5.0-11.0 (optimum, pH 7.0). The major menaquinone was MK-8(H 4 ) and the major cellular fatty acids (>10 %) were identified as iso-C 16 : 0 and C 17 : 1  ω 8 c . Strain YIM 123512 T had ll-2, 6-diaminopimelic acid as the diagnostic cell-wall diamino acid. The major polar lipids were found to be diphosphatidylglycerol, phosphatidylglycerol, two unknown phospholipids and one unknown lipid. The genomic DNA G+C content of strain YIM 123512 T was 72.7 mol% based on its draft genome sequence. Phylogenetic analysis based on the 16S rRNA gene sequence indicated that strain YIM 123512 T appeared to be most closely related to Nocardioides halotolerans KCTC 19274 T (97.7 % sequence similarity) and to belong to the genus Nocardioides . The sequence similarity values of strain YIM 123512 T to other currently described type strains of the genus Nocardioides were less than 97.0 %. Based on the draft genome sequence, the novel strain showed an average nucleotide identity value of 80.2 % and a digital DNA-DNA hybridization value of 23.1 % with the reference strain N. halotolerans KCTC 19274 T . The results of the polyphasic taxonomic study including phenotypic, chemotaxonomic and phylogenetic analyses show that strain YIM 123512 T represents a novel species of the genus Nocardioides , for which the name Nocardioides flavescens sp. nov. is proposed. The type strain is YIM 123512 T (=KCTC 49303 T =CGMCC 4.7628 T ).