Rhodoalgimonas zhirmunskyi gen. nov., sp. nov., a Marine Alphaproteobacterium Isolated from the Pacific Red Alga Ahnfeltia tobuchiensis: Phenotypic Characterization and Pan-Genome Analysis

A novel Gram-staining negative, strictly aerobic, rod-shaped, and non-motile bacterium, designated strain 10Alg 79T, was isolated from the red alga Ahnfeltia tobuchiensis. A phylogenetic analysis based on 16S rRNA gene sequences placed the novel strain within the family Roseobacteraceae, class Alphaproteobacteria, phylum Pseudomonadota, where the nearest neighbor was Shimia sediminis ZQ172T (97.33% of identity). However, a phylogenomic study clearly showed that strain 10Alg 79T forms a distinct evolutionary lineage at the genus level within the family Roseobacteraceae combining with strains Aquicoccus porphyridii L1 8-17T, Marimonas arenosa KCTC 52189T, and Lentibacter algarum DSM 24677T. The ANI, AAI, and dDDH values between them were 75.63–78.15%, 67.41–73.08%, and 18.8–19.8%, respectively. The genome comprises 3,754,741 bp with a DNA GC content of 62.1 mol%. The prevalent fatty acids of strain 10Alg 79T were C18:1 ω7c and C16:0. The polar lipid profile consisted of phosphatidylethanolamine, phosphatidylglycerol, phosphatidylcholine, an unidentified aminolipid, an unidentified phospholipid and an unidentified lipid. A pan-genome analysis showed that the unique part of the 10Alg 79T genome consists of 13 genus-specific clusters and 413 singletons. The annotated singletons were more often related to transport protein systems, transcriptional regulators, and enzymes. A functional annotation of the draft genome sequence revealed that this bacterium could be a source of a new phosphorylase, which may be used for phosphoglycoside synthesis. A combination of the genotypic and phenotypic data showed that the bacterial isolate represents a novel species and a novel genus, for which the name Rhodoalgimonas zhirmunskyi gen. nov., sp. nov. is proposed. The type strain is 10Alg 79T (=KCTC 72611T = KMM 6723T).


Introduction
Red algae are widely distributed in marine environments and tend to be colonized by rich and diverse communities of bacteria, some of which may be cultured under lab-oratory conditions [1][2][3][4].Many novel bacterial species associated with different red algae have been described in recent years.The strains belonging to a new genus and species Algibacillus agarilyticus (the family Alteromonadaceae, class Gammaproteobacteria), and two new species Marinomonas algarum (the family Oceanospirillaceae, class Gammaproteobacteria) and Maribacter algarum (the family Flavobacteriaceae, class Flavobacteriia) were isolated from the red alga Gelidium amansii collected from coastal waters in Wenhai (Shandong province, China) [5][6][7].The novel members of the genera Marinomonas and Maribacter, Marinomonas agarivorans and Maribacter algicola, were also recovered from the surfaces of the red algae Gracillaria bladgettii and Porphyridium marinum, respectively [8,9].The representatives of three new species of the family Flavobacteriaceae, Muricauda (formerly Flagellimonas) algicola, Psychroserpens luteolus, and Tenacibaculum aquimarinum, were described as epiphytic bacteria of the red algae Asparagopsis taxiformis, Gelidium sp., and Chondrus sp.collected from Korean and Chinese coastal waters [10][11][12].The new species Thalassotalea algicola (family Colwelliaceae, class Gammaproteobacteria) was isolated from a red alga Porphyra sp.collected from the Chinese coast in Weihai [13].The new genus and species Hwanghaeella grinnelliae (the family Rhodospirillaceae, class Alphaproteobacteria) was isolated from a red marine alga Grinnellia sp. in the Yellow Sea of the Republic of Korea [14].Other representatives of the class Alphaproteobacteria associated with the Pacific red alga Polysiphonia sp. were placed as new species of the family Roseobacteraceae (formerly Rhodobacteraceae), Roseibium (formerly Labrenzia) polysiphoniae [15].The new species Aquimarina algiphila (the family Flavobacteriaceae, class Flavobacteriia), and the new genus and species Algicella marina (the family Paracoccaceae, class Alphaproteobacteria) were recovered from a common inhabitant of the Sea of Japan, Tichocarpus crinitus [16,17].Three new species of the family Flavobacteriaceae, Aureibaculum algae, Flavobacterium ahnfeltiae, and Polaribacter staleyi, were isolated from the red alga Ahnfeltia tobuchiensis collected from coastal waters of the Okhotsk Sea [17][18][19].
The aim of this report is to provide a genomic characterization and taxonomic description of another associate of the red alga Ahnfeltia tobuchiensis, designated as strain 10Alg 79 T (KMM 6723 T ).This isolate is proposed as a new member of the family Roseobacteraceae (formerly Rhodobacteraceae) based on the study of its phenotypic, chemotaxonomic, and genomic characteristics.

Bacterial Isolation and Maintenance
Strain 10Alg 79 T was isolated from the red alga Ahnfeltia tobuchiensis collected near Island Paramushir, Kuril Isls, Okhotsk Sea, Russia, using a standard dilution plating method.The sample of algal fronds (5 g) was homogenized in 10 mL sterile seawater in a glass homogenizer, and 0.1 mL of homogenate was spread onto Marine Agar 2216 (MA, Difco) plates.The novel isolates were obtained from a single colony after incubation of the plate at 28 • C for 7 days.After primary isolation and purification, the strains were cultivated at 28 • C on the same medium and stored at −80 • C in marine broth (Difco, Sparks, MD, USA) supplemented with 20% (v/v) glycerol.The type strains Aquicoccus porphyridii JCM 31543 T and Marimonas arenosa KCTC 52189 T obtained from the Japan Collection of Microorganisms (JCM) and the Korean Collection for Type Cultures (KCTC), respectively, were used as reference strains in the parallel tests for this study.The data for Lentibacter algarum ZXM100 T neighboring to the new isolate on the genomic tree were also included in all tables as a reference strain.
Fatty acid methyl esters and polar lipids of strain 10Alg 79 T and its closest phylogenetic relatives, Aquicoccus porphyridii JCM 31543 T and Marimonas arenosa KCTC 52189 T , were extracted and analyzed as described previously [23] using cells grown on MA for 48 h at 28 • C. Isoprenoid quinones were extracted with chloroform/methanol (2:1, v/v) and purified with TLC using a mixture of n-hexane and diethyl ether (85:15, v/v) as the solvent.Isoprenoid quinone composition was characterized with HPLC (Shimadzu LC-10A, Shimadzu, Kyoto, Japan) using a reversed phase type Supelcosil LC-18 column (15 cm × 4.6 mm) and acetonitrile/2-propanol (65:35, v/v) as a mobile phase at a flow rate of 0.5 mL min −1 , as described previously [24].The column was kept at 40 • C. Quinones were detected by monitoring at 275 nm.
The analysis of 16S rRNA gene sequences was performed on the EzBioCloud server [26], and the phylogenetic analysis was conducted using MEGA X software, version 10.2.1 [27], with the neighbor-joining (NJ), maximum-likelihood (ML), and maximum-parsimony (MP) methods.Genetic distances were calculated according to the Kimura two-parameter model [28], and bootstrap values were obtained from 500-1000 alternative trees.
The maximum-likelihood phylogeny of RpoC sequences translated from genome sequences was calculated using the IQ-TREE web server, version 1.6.12[29], with 100 nonparametric bootstrap replicates and the LG+I+G4 substitution model determined using ModelFinder [30].

Whole-Genome Sequencing and Genome-Based Phylogenetic Analysis
The genomic DNA of strain 10Alg 79 T extracted as described above in Section 2.4 was verified using DNA gel electrophoresis and quantified in a Qubit 4.0 Fluorometer (Thermo Fisher Scientific, Waltham, MA, USA).The DNA sequencing library was prepared using Nextera DNA Flex kits (Illumina, San Diego, CA, USA) and was sequenced using pairedend runs on an Illumina MiSeq platform with a 150 bp read length.The row reads were trimmed using Trimmomatic, version 0.39 [31], and their quality was assessed using FastQC, version 0.11.8 (https://www.bioinformatics.babraham.ac.uk/projects/fastqc/, accessed on 21 August 2021).Filtered reads were assembled into contigs with SPAdes, version 3. 15.3 [32].The quality of assembly was analyzed using QUAST, version 5.0.2 [33].Genome completeness and contamination were estimated using CheckM, version 1.1.3,based on the taxonomic-specific workflow (family Rhodobacteriaceae) [34].The genome was annotated using the NCBI Prokaryotic Genome Annotation Pipeline (PGAP), Rapid Annotation using Subsystem Technology (RAST), and the Pathosystems Resource Integration Center (PATRIC) [35][36][37].Comparisons of the average nucleotide identity (ANI), average amino acid identity (AAI), and in silico DNA-DNA hybridization (dDDH) values of strain 10Alg 79 T and its closest neighbors were performed with the online server ANI/AAI-Matrix [38] and TYGS platform [39], respectively.
To identify carbohydrate-active enzymes (CAZymes), the dbCAN2 meta server, version 11, was used with default settings (http://cys.bios.niu.edu/dbCAN2,accessed on 1 December 2022) [43].Predictions using two of the three algorithms integrated within the server (DIAMOND, HMMER, and dbCAN-sub) were considered sufficient for CAZy family assignments.The relative abundances of CAZymess were visualized with heat maps using pheatmap, version 1.0.12, in RStudio, version 2022.02.0+443, with R, version 4.1.3.Annotation of secondary metabolite biosynthetic gene clusters was conducted using antiSMASH server, version 7.0.0beta1-67b538a9(https://antismash.secondarymetabolites.org/#!/start, accessed on 12 December 2022) [44].The genomic regions containing NRPS-like fragment and Type I PKS (Polyketide synthase) biosynthetic gene clusters were extracted from GBK files of the genomes using Geneious Pro software, version 4.8 [45].Generated GBK files were modified by adding custom color feature qualifiers, according to antiSMASH conventional coloring.Pairwise comparisons of each locus between four genomes were carried out using BLASTn (BLAST version 2.11.0+) run in EasyFig (version 2.2.5) [46].Synteny plots were visualized using Easyfig with a minimum of 100 bp BLAST hits.Fonts and sizes in all figures were edited manually in Adobe Photoshop CC 2018 for better visualization.
The draft genome of M. arenosa KCTC 52189 T was obtained in this study as described above for the draft genome of 10Alg 79 T and used in the comparative genome analysis.
Therefore, the position of strain 10Alg 79 T was further determined using rpoC gene sequences extracted from type strain genomes of genera affiliated with the family Roseobacteraceae (formerly Rhodobacteraceae).RpoC protein sequences have been shown to be suitable for the Roseobacter group phylogeny [17,47].An ML phylogenetic tree inferred on 43 RpoC protein sequences showed that 10Alg 79 T forms a poorly supported clade together with M. arenosa KCTC 52189 T [48], A. porphyridii L1 8-17 T [49], and L. algarum DSM 24677 T [50] (Figure 2).
These values were significantly below the boundaries for the demarcation of bacterial species [51,52].The AAI values between 10Alg 79 T and the representatives were 67.41-73.08%,which did not exceed the declared genus boundaries of 80% [53].Thus, the phylogenomic analysis based on the genome relatedness indexes clearly supported 10Alg 79 T as a novel type species of a new genus in the family Rosebacteraceae (formerly Rhodobacteraceae).
Remarkably, a search for 16S rRNA sequences homologous to 10Alg 79 T in NCBI, Greengenes, EzBioCloud, and Silva databases revealed only two 16S rRNAs with 99% identity (JQ215453.1 and JQ212029.1).Both originated from uncultured bacterium clones isolated from the dolphin Tursiops truncates in 2011 and 2013.This means that 10Alg 79 T represents a new species of the rare genus of the family Roseobacteraceae.These values were significantly below the boundaries for the demarcation of bacterial species [51,52].The AAI values between 10Alg 79 T and the representatives were 67.41-73.08%,which did not exceed the declared genus boundaries of 80% [53].Thus, the phylogenomic analysis based on the genome relatedness indexes clearly supported 10Alg 79 T as a novel type species of a new genus in the family Rosebacteraceae (formerly Rhodobacteraceae).Remarkably, a search for 16S rRNA sequences homologous to 10Alg 79 T in NCBI, Greengenes, EzBioCloud, and Silva databases revealed only two 16S rRNAs with 99% identity (JQ215453.1 and JQ212029.1).Both originated from uncultured bacterium clones isolated from the dolphin Tursiops truncates in 2011 and 2013.This means that 10Alg 79 T represents a new species of the rare genus of the family Roseobacteraceae.

Genomic Characteristics and Analysis of Genus-Related Features
The draft genome of strain 10Alg 79 T was de novo assembled into 73 contigs, with an N50 value of 702,161 bp and an L50 value of two.The genome size was estimated at 3,754,741 bp in length with an overall GC content of 62.1%.The genome-extracted 16S rRNA gene sequence was 100% identical to the PCR-amplified one.The genome contains 3739 coding sequences, 41 tRNAs, and 4 rRNA genes (one each of 16S and 23S and two genes of 5S).A comparison between genome features of 10Alg 79 T and the type strains of closely related genera is presented in Table 1.These characteristics satisfy the proposed minimal standards for bacterial taxonomy [54].

Genomic Characteristics and Analysis of Genus-Related Features
The draft genome of strain 10Alg 79 T was de novo assembled into 73 contigs, with an N 50 value of 702,161 bp and an L 50 value of two.The genome size was estimated at 3,754,741 bp in length with an overall GC content of 62.1%.The genome-extracted 16S rRNA gene sequence was 100% identical to the PCR-amplified one.The genome contains 3739 coding sequences, 41 tRNAs, and 4 rRNA genes (one each of 16S and 23S and two genes of 5S).A comparison between genome features of 10Alg 79 T and the type strains of closely related genera is presented in Table 1.These characteristics satisfy the proposed minimal standards for bacterial taxonomy [54].
To clarify genus-related features, a pan-genome analysis of 10Alg 79 T and representatives of three closely related genera (M.arenosa KCTC 52189 T , A. porphyridii L1 8-17 T , and L. algarum DSM 24677 T ) was performed using orthologous clustering with the OrthoVenn2 and OrthoVenn3 servers [41,42].The analysis revealed 3374 orthologous gene clusters (gene families); 1602 single-copy clusters consisted of only single-copy orthologs presented in all lineages of the clade.The core and accessory (without unique genes) genomes consisted of 2167 and 1022 orthologous clusters, respectively (Figure 4).Most of the 10Alg 79 T clusters were shared with M. arenosa KCTC 52189 T (141) and A. porphyridii L1 8-17 T (116) (Figure 4).The results are in good agreement with the taxonomic position of 10Alg 79 T on the genomic tree (Figure 3); M. arenosa KCTC 52189 T and A. porphyridii L1 8-17 T were more related to 10Alg 79 T than L. algarum DSM 24677 T .A total of 185 were identified as genus-specific clusters (Figure 4).To predict metabolism of representatives of the nearest genera, genomes were first screened for the presence of genes of central and peripheral carbon metabolism due to the ability to grow on various carbon resources (Table 2).Based on the KEGG annotation, the genomes possess almost all genes of the glycolysis/gluconeogenesis pathways except for a key gene encoding 6-phosphofructokinase (EC 2.7.1.11) in the Embden-Meyerhof pathway.However, a modified pathway might incorporate the pentose phosphate pathway (PPP) due to genes encoding transaldolase (EC 2.2.1.2) and transketolase (EC 2.2.1.1).Moreover, genes found for the Entner-Doudoroff pathway (EDP), excluding L. algarum DSM 24677 T , might encode the metabolism of glucose-6-phosphate into glyceraldehyde-3-phosphate and pyruvate.All the genomes were devoid of pgd encoding 6-phosphogluconate dehydrogenase (EC 1.1.1.44)in the oxidative phase of PPP.Semi-phosphorylative and non-phosphorilative EDP were presented by a partial set of genes.This means that the species are not able to utilize glucose through these canonical pathways and produce NADPH using PPP.All the genomes encode a complete tricarboxylic acid (TCA) cycle.Interestingly, M. arenosa KCTC 52189 T also encodes a complete Calvin-Benson-Bassham cycle, in which a key enzyme RuBisCO (EC 4.1.1.39),is encoded by rbcLS and accompanied by molecular chaperon genes cbbQ and ccbO to enhance CO 2 fixation activity [55].
Genome mining of 10Alg 79 T for secondary metabolite biosynthetic gene clusters (BGS) revealed five biotechnologically significant BGSs for the putative synthesis of non-ribosomal peptide synthetase (NRPS)-like and polyketide synthetase (PKS/NRPS hybrid), ribosomally synthesized and post-translationally modified peptides (RiPP-like, with methanobactin-like DUF692 domain), two homoserine lactones (hserlactone), and one important osmoprotectant ectoine.The most of these BGSs were also characteristic for the closely related genera Marimonas, Lentibacter, and Aquicoccus.For example, 10Alg 79 T sheared up to 90% of genes encoding the PKS/NRPS hybrid system with 50-88% of gene identities (Figure 6), as estimated by [44].
Out of 16,163 genes identified in the clade pan-genome, 2312 genes were assigned as singletons without any orthologues among the compared genomes.Among them 413 singletons were found in the 10Alg 79 T genome, followed by L. algarum DSM 24677 T (410), A. porphyridii L1 8-17 T (713), and M. arenosa KCTC 52189 T (775).An analysis of a unique part of each genome showed that most were annotated as hypothetical proteins (more than 60%).The annotated singletons were more often related to transport protein systems, transcriptional regulators, and enzymes.
For comparison, the M. arenosa KCTC 52189 T genome represents five orthologous genes and one singleton gene for alkaline phosphatases, followed by L. algarum DSM 24677 T with only two orthologous genes and A. porphyridii L1 8-17 T with only one orthologous gene.Therefore, attention should be paid to nitrogen metabolisms due to the number of singleton genes belonging to these subsystems.Remarkably, the 10Alg 79 T genome contains two urease operons for urea catabolism; the first ureDABCEFG was shared with those of the other three genomes, and the second ureEFACGD was composed of singleton genes homologous to one of the numerous urease operons of Salipiger spp.(about 80% amino acid identity).This second ure operon was accompanied by an additional urea transport operon utrABCDE, which is like that in Salipiger spp, indicating past lateral gene transfer events.Additionally, 10Alg79 harbors singletons encoding N-methylhydantoinase A and B (EC 3.5.2.14), involved in the metabolism of nitrogen-containing compounds.

Morphological, Physiological, and Biochemical Characteristics
Strain 10Alg 79 T has many properties in common with its closest relatives.They were found to be strictly aerobic, heterotrophic, and non-motile rods that can produce alkaline phosphatase, esterase (C4), esterase lipase (C8), leucine arylamidase, acid phosphatase, naphthol-AS-BI-phosphohydrolase, and catalase and oxidase but not lipase (C14), cystine arylamidase, trypsin, α-chymotrypsin, α-galactosidase, β-galactosidase, β-glucuronidase, α-glucosidase, β-glucosidase, N-acetyl-β-glucosaminidase, α-mannosidase, α-fucosidase, agarase, or caseinase.The novel isolate was distinguished from all neighbors by the minimal temperature for growth and amylase production.The sets of phenotypic traits including the ability to grow without NaCl or seawater, to reduce nitrate to nitrite, to produce acetoin and hydrogen sulfide, to hydrolyze aesculun, gelatin, tyrosine, and Tweens 40 and 80, taken together with the ability to utilize various carbohydrates, can help to discriminate strain 10Alg 79 T from each of its relatives (Table 2).

Chemotaxonomy
The fatty acid composition of strain 10Alg 79 T and reference strains A. porphyridii JCM 31543 T and M. arenosa KCTC 52189 T are listed in Table 3.The predominant fatty acids of strain 10Alg 79 T were C18:1 ω7c (82.1%) and C16:0 (8.2%) (Table 3).The fatty acid composition of the strain studied was similar to that of the reference strains, although differences were found in the proportions of some fatty acids (Table 3).The polar lipid profile of strain 10Alg 79 T contained phosphatidylethanolamine, phosphatidylglycerol, phosphatidylcholine, an unidentified phospholipid, an unidentified aminolipid, and an unidentified lipid and was consistent with that of the reference species (Supplementary Figure S1; Table 3).However, the presence of an unidentified phospholipid distinguished the bacterial isolate from M. arenosa KCTC 52189 T and L. algarum ZXM100 T .Moreover, strain 10Alg 79 T can be differentiated from M. arenosa KCTC 52189 T by the absence of an unknown glycolipid (Table 3).The predominant respiratory quinone was Q-10.

Microorganisms 2023 , 17 Figure 1 .
Figure 1.ML 16S rRNA tree showing phylogenetic relationships between the novel strain 10Alg 79 T (KMM 6723 T ) and members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values shown as percentage are based on 500 replicates.Bars are 0.5 substitutions per nucleotide.Strain Caulobacter vibrioides CB51 T /DSM 9893 T was used as an outgroup.GenBank/EMBL/DDB accession numbers are given in parentheses.

Figure 1 .
Figure 1.ML 16S rRNA tree showing phylogenetic relationships between the novel strain 10Alg 79 T (KMM 6723 T ) and members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values shown as percentage are based on 500 replicates.Bars are 0.5 substitutions per nucleotide.Strain Caulobacter vibrioides CB51 T /DSM 9893 T was used as an outgroup.GenBank/EMBL/DDB accession numbers are given in parentheses.

Figure 2 .
Figure 2. ML RpoC tree showing phylogenetic relationships between the novel strain 10Alg 79 T (KMM 6723 T ) and 43 members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values are based on 500 replicates and shown as percentages greater than 50.Bars are 0.5 substitutions per amino acid position.Strain Caulobacter vibrioides DSM 9893 T was used as an outgroup.Ge-Bank/EMBL/DDB accession numbers are given in parentheses.

Figure 2 .
Figure 2. ML RpoC tree showing phylogenetic relationships between the novel strain 10Alg 79 T (KMM 6723 T ) and 43 members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values are based on 500 replicates and shown as percentages greater than 50.Bars are 0.5 substitutions per amino acid position.Strain Caulobacter vibrioides DSM 9893 T was used as an outgroup.GeBank/EMBL/DDB accession numbers are given in parentheses.

Figure 3 .
Figure 3. ML genomic tree showing phylogenetic relationships between the strain 10Alg 79 T (KMM 6723 T ) and members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values shown as percentages greater than 50 are based on 100 replicates.Bar is 0.2 substitutions per amino acid position.Strain Caulobacter vibrioides DSM 9893 T was used as an outgroup.Gen-Bank/EMBL/DDB accession numbers are given in parentheses.

Figure 3 .
Figure 3. ML genomic tree showing phylogenetic relationships between the strain 10Alg 79 T (KMM 6723 T ) and members of the family Rosebacteraceae (formerly Rhodobacteriaceae).Bootstrap values shown as percentages greater than 50 are based on 100 replicates.Bar is 0.2 substitutions per amino acid position.Strain Caulobacter vibrioides DSM 9893 T was used as an outgroup.GenBank/EMBL/DDB accession numbers are given in parentheses.

Figure 4 .
Figure 4. Venn diagram showing the distribution of shared orthologous clusters among genomes of 10Alg 79 T , A. porphyridii L1 8-17 T , M. arenosa KCTC 52189 T , and L. algarum DSM 24677 T .The orthologous clusters are comprised of proteins predicted using RAST [36].The number of orthologous clusters for each genome is shown in vertical bar chart.

Figure 5 .
Figure 5.The distribution of CAZymes across genomes of KMM 6723 T (=10Alg 79 T ) and close relatives.The heat maps show the number of genes assigned to individual CAZyme families.Rows are clustered using Euclidean distances.GH, glycoside hydrolases; GT, glycosyltransferases, CE, carbohydrate esterase; AA, auxiliary activity; CBM, carbohydrate-binding module.

Figure 5 .
Figure 5.The distribution of CAZymes across genomes of KMM 6723 T (=10Alg 79 T ) and close relatives.The heat maps show the number of genes assigned to individual CAZyme families.Rows are clustered using Euclidean distances.GH, glycoside hydrolases; GT, glycosyltransferases, CE, carbohydrate esterase; AA, auxiliary activity; CBM, carbohydrate-binding module.Microorganisms 2023, 11, x FOR PEER REVIEW 12 of 17

Figure 6 .
Figure 6.Comparative analysis of synteny between NRPS-like fragment and Type I PKS BGCs in the genomes of strain 10Alg 79 T (KMM 6723 T ) and close relatives.Genes are colored based on their annotation as indicated at the bottom.The locus regions are NOI20_03420 to NOI20_03580 in the KMM 6723 T genome, NO357_05410 to NO357_05560 in the M. arenosa KCTC 52189 T genome, BLV93_RS04080 to BLV93_RS03920 in the L. algarum DSM 24677 T genome, and FLO80_RS04395 to FLO80_RS04560 in the A. porphyridii JCM 31543 T genome.

Figure 6 .
Figure 6.Comparative analysis of synteny between NRPS-like fragment and Type I PKS BGCs in the genomes of strain 10Alg 79 T (KMM 6723 T ) and close relatives.Genes are colored based on their annotation as indicated at the bottom.The locus regions are NOI20_03420 to NOI20_03580 in the KMM 6723 T genome, NO357_05410 to NO357_05560 in the M. arenosa KCTC 52189 T genome, BLV93_RS04080 to BLV93_RS03920 in the L. algarum DSM 24677 T genome, and FLO80_RS04395 to FLO80_RS04560 in the A. porphyridii JCM 31543 T genome.

Table 1 .
Genomic features of 10Alg 79 T and three type strains of closely related genera.

Table 2 .
Phenotypic characteristics differentiating strain 10Alg 79 T and related genera of the family Roseobacteraceae.