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Reclassification of 11 Members of the Family Rhodobacteraceae at Genus and Species Levels and Proposal of Pseudogemmobacter hezensis sp. nov.

A novel Gram-stain-negative, aerobic, motile bacterial strain, D13-10-4-6T, was isolated from the bark sample of Populus × euramericana. The strain could grow at 15–35°C, at pH 6–10 and in 0–4% (w/v) NaCl, and the strain tested positive for oxidase and catalase activities. The main polar lipids were phosphatidylmonomethylethanolamine, diphosphatidylglycerol, phosphatidylglycerol, and phosphatidylethanolamine. The main respiratory quinone was Q-10, and the predominant fatty acid was C18:1 ω7c. The phylogenetic analyses showed that the strain belonged to the genus Pseudogemmobacter of the family Rhodobacteraceae. The family Rhodobacteraceae is an ecologically diverse group that includes bacteria from aquatic to terrestrial ecosystems. As a consequence, the classification of the family Rhodobacteraceae is difficult, not least when the early taxonomy work relied heavily on 16S rRNA gene analysis. Recently, the taxonomic status of many members of the family has been revised based on the genome analysis; however, there are still some classification conflicts due to the lack of genome sequences and parallel publication time. In this study, phylogenetic trees based on 16S rRNA gene, gyrB gene, and 120 concatenated proteins, the average amino acid identity (AAI) and percentage of conserved proteins (POCP) have been used for the analysis of strain D13-10-4-6T and other members of 15 genera within the family to further clarify their taxonomic relationships. For the data of phylogeny, AAI, and POCP, the taxonomic proposals are (1) reclassification of Rhodobacter tardus as the type species of a novel genus, Stagnihabitans gen. nov., as Stagnihabitans tardus comb. nov.; (2) reclassification of Tabrizicola alkalilacus, Tabrizicola sediminis, Tabrizicola algicola into a novel genus, Pseudotabrizicola gen. nov., as Pseudotabrizicola alkalilacus comb. nov., Pseudotabrizicola sediminis comb. nov., Pseudotabrizicola algicola comb. nov.; (3) reclassification of Rhodobacter sediminicola into the genus Cereibacter as Cereibacter sediminicola comb. nov.; (4) reclassification of Rhodobacter flagellatus, Rhodobacter thermarum, and Xinfangfangia soli into the genus Tabrizicola as Tabrizicola flagellatus comb. nov., Tabrizicola thermarum comb. Nov., and Tabrizicola soli comb. nov.; (5) reclassification of Xinfangfangia humi into the genus Pseudogemmobacter as Pseudogemmobacter humicola comb. nov.; (6) classification of strain D13-10-4-6T as a novel species of the genus Pseudogemmobacter, for which the name P. hezensis sp. nov. is proposed, the type strain is D13-10-4-6T (= CFCC 12033T = KCTC 82215T).

A novel Gram-stain-negative, aerobic, motile bacterial strain, D13-10-4-6 T , was isolated from the bark sample of Populus × euramericana. The strain could grow at 15-35 • C, at pH 6-10 and in 0-4% (w/v) NaCl, and the strain tested positive for oxidase and catalase activities. The main polar lipids were phosphatidylmonomethylethanolamine, diphosphatidylglycerol, phosphatidylglycerol, and phosphatidylethanolamine. The main respiratory quinone was Q-10, and the predominant fatty acid was C 18:1 ω7c. The phylogenetic analyses showed that the strain belonged to the genus Pseudogemmobacter of the family Rhodobacteraceae. The family Rhodobacteraceae is an ecologically diverse group that includes bacteria from aquatic to terrestrial ecosystems. As a consequence, the classification of the family Rhodobacteraceae is difficult, not least when the early taxonomy work relied heavily on 16S rRNA gene analysis. Recently, the taxonomic status of many members of the family has been revised based on the genome analysis; however, there are still some classification conflicts due to the lack of genome sequences and parallel publication time. In this study, phylogenetic trees based on 16S rRNA gene, gyrB gene, and 120 concatenated proteins, the average amino acid identity (AAI) and percentage of conserved proteins (POCP) have been used for the analysis of strain D13-10-4-6 T and other members of 15 genera within the family to further clarify their taxonomic relationships. For the data of phylogeny, AAI, and POCP, the taxonomic proposals are (1) reclassification of Rhodobacter tardus as the type species of a novel genus, Stagnihabitans gen. nov., as Stagnihabitans tardus comb. nov.; (2) reclassification of Tabrizicola alkalilacus, Tabrizicola sediminis, Tabrizicola algicola into a novel genus, Pseudotabrizicola gen. nov., as Pseudotabrizicola alkalilacus comb. nov., Pseudotabrizicola sediminis comb. nov., Pseudotabrizicola algicola comb. nov.; (3) reclassification of Rhodobacter sediminicola into the genus Cereibacter as Cereibacter sediminicola comb. nov.; (4) reclassification of Rhodobacter flagellatus, Rhodobacter thermarum, and Xinfangfangia soli into the genus Tabrizicola as Tabrizicola flagellatus comb. nov., Tabrizicola thermarum comb. Nov., and

INTRODUCTION
Populus × euramericana cakers on poplar trees were found in China for many years, and the stem or branch bark of the diseased tree was cracked and exuded frothy fluid. During our investigation of the bacterial diversity in Populus × euramericana caker, strain D13-10-4-6 T was isolated from the symptomatic bark of Populus × euramericana caker. The phylogenetic analyses showed that the strain belonged to the genus Pseudogemmobacter of the family Rhodobacteraceae. The family Rhodobacteraceae, described by Garrity et al. (2005), the so-called purple nonsulfur bacteria (Imhoff et al., 1998), is one of the major subdivisions of the class Alphaproteobacteria. It is ecologically and phenotypically diverse, and most of the members of the family have been found in various marine environments, including seawater, sea sediments, sea ice, coastal biofilms, marine animal tissues, and seaweeds (Selje et al., 2004;Buchan et al., 2005;Brinkhoff et al., 2008). At the time of writing, the family included more than 180 genera with validated names. 1 The early classification of the genera within the family Rhodobacteraceae relied heavily on the analysis of 16S rRNA gene sequence and resulted in several non-monophyletic genera, for instance, the genus Rhodobacter (Imhoff et al., 1984). The genus Rhodobacter was reclassified by Suresh et al. (2019) and Hördt et al. (2020) based on the genome analysis. The members of the genus were divided into five distinct clades in the 16S rRNA gene-based phylogenetic tree constructed by Suresh et al. (2019). The Cereibacter sphaeroides (formerly Rhodobacter sphaeroides) clade was reclassified into the genus Luteovulum (Suresh et al., 2019) and then transferred into the genus Cereibacter by Hördt et al. (2020). However, Rhodobacter alkalitolerans, which belongs to the C. sphaeroides clade, was not reclassified due to the lack of genomic sequence. At present, the genus Rhodobacter contains 13 species with validated names according to the List of Prokaryotic names with Standing in Nomenclature (LSPN). 2 The taxonomic status of the recently described Rhodobacter species, R. thermarum (Khan et al., 2019), R. flagellatus (Xian et al., 2020), R. sediminicola (Suresh et al., 2020), and R. tardus (Sheu et al., 2020), was also not included in the early reclassification of Rhodobacter due to the nearly parallel time of description, causing confusion in the classification of the genus Rhodobacter.
Along with the advances in whole-genome sequencing technologies, several methods for taxonomic classification at the species and genus levels have been developed. The new standards for species recognition are developed using digital whole-genome comparisons, such as average nucleotide identities (ANIs) (Konstantinidis and Tiedje, 2005) and genome-togenome-distance calculations (GGDCs) (Richter and Rosselló-Móra, 2009;Meier-Kolthoff et al., 2013). The average amino acid identity (AAI) (Luo et al., 2014;Rodriguez-R and Konstantinidis, 2014) and percentage of conserved proteins (POCPs) (Qin et al., 2014), which are methods of measuring amino acidlevel genomic similarity between protein-coding regions, have been used in the delineation of prokaryotic organisms at the genus level. Furthermore, the phylogenetic analysis based on the whole-genome sequence has been recently encouraged for the taxonomy of prokaryotes owing to its robustness and repeatability (Chun et al., 2018). Nowadays, along with those methods, the reclassification of prokaryotes at class (Hördt et al., 2020), order (Orata et al., 2018), family (Liang et al., 2021), and genus (Suresh et al., 2019) levels has been done.
In this study, phylogenetic analysis based on the 16S rRNA gene, gyrB gene, and genomes sequence, as well as ANI, AAI, and POCP, was used to confirm the taxonomic relationship of the novel strain and its closely related members (e.g., members of the genus Xinfangfangia and Rhodobacter) in the family Rhodobacteraceae.

Strain and Culture Conditions
Strain D13-10-4-6 T was isolated from the bark sample of Populus × euramericana collected from Heze, Shandong Province, China (34 • 82 N, 115 • 46 E) as previously described (Li et al., 2015). In brief, the bark samples were sterilized for 30 s with 70% ethanol, and then exposed to 4% (v/v) sodium hypochlorite for 3 min. After rinsing in sterile water three times, the samples with 2 ml sterile water were transferred to sterile mortar and ground with a pestle, respectively. The obtained solution was then shaken for 30 min at 30 • C. The suspension with a dilution series was spread on tryptic soy agar (TSA, Difco). After 2 days of incubation on TSA plates at 30 • C, single colonies were selected and cultured on a new plate, and were then preserved at -80 • C with a supplement of 20% (v/v) glycerol.

Phylogenetic Analysis
The 16S rRNA gene of strain D13-10-4-6 T was amplified by the primers 27F/1492R (Lane, 1991). The similarity of the 16S rRNA gene sequence between the strain D13-10-4-6 T and the validly published bacterial species was determined using EzBio-Cloud's identify service 3 (Yoon et al., 2017). The 16S rRNA gene sequence of the related strains was obtained from GenBank for the phylogenetic analysis. After multiple sequence alignment with Clustal W, the phylogenetic analysis was carried out using MEGA X by the neighbor-joining, maximumlikelihood, and maximum-parsimony methods (Kumar et al., 2018). Aquidulcibacter paucihalophilus TH1-2 T was used as an outgroup. The phylogenetic trees were evaluated by 1,000 bootstrap resamplings.
The gyrB gene sequences of the strain D13-10-4-6 T were obtained from its genomic sequences according to Altschul et al. (1990), and a 1,050 bp sequence was obtained. The gyrB gene sequences of the related strains were obtained from GenBank or their genome sequences. The phylogenetic trees based on the gyrB gene sequence were constructed using the maximumlikelihood, neighbor-joining, and maximum-parsimony methods as a description of 16S rRNA gene phylogenetic analysis.
Concatenated protein tree has a higher recognition than single phylogenetic marker gene tree (e.g., 16S rRNA and gyrB) for bacterial taxonomy (Ciccarelli et al., 2006;Thiergart et al., 2014), and has been widely used in solving bacterial taxonomy (Hördt et al., 2020;Xu et al., 2021). The genome sequences of the strain D13-10-4-6 T and its related strains retrieved from GenBank (Supplementary Table 1) were used to construct the phylogenetic tree. A concatenated alignment of 120 ubiquitous single-copy proteins of the related strains was performed by GTDB-Tk v1.5.1 4 using the classify_wf command (Chaumeil et al., 2020). The alignment file was used to construct the maximum-likelihood tree with IQ-TREE 2.1.4-beta (Minh et al., 2020), and the best model was automatically selected by ModerFinder for the ML tree. The tree was visualized and edited with iTOL (Letunic and Bork, 2021).

Phylogenomic Metric Calculations
Average nucleotide identity (ANI is a measure of similarity between two genomic sequences, which is a useful tool to differentiate bacterial species in common with DNA-DNA hybridization (DDH) (Goris et al., 2007;Richter and Rosselló-Móra, 2009). The ANI values among the novel strain D13-10-4-6 T and its closely related reference strains (P. bohemicus Cd-10 T , X. humi IMT-291 T ) were determined using OrthoANI (Lee et al., 2016). The GGDC 5 was used to calculate the dDDH values among the novel and its closely related reference strains (Meier-Kolthoff et al., 2013). The analysis of the average AAI and POCP among the strains in this work was carried out with CompareM 6 and a Python script (POCP) 7 (Xu et al., 2020), respectively. The pangenome analysis was carried out with BPGA (Chaudhari et al., 2016) with default parameters.

Chemotaxonomic Characterization
The strain D13-10-4-6 T was shaken for 48 h in a tryptic soy broth (TSB; Difco) at 30 • C, then collected by centrifuging at 10,000 rpm for 4 min. The harvested cells were freeze-dried and used to analyze the polar lipid and respiratory quinone. Polar lipids were analyzed by two-dimensional thin-layer chromatography as described by Minnikin et al. (1984). Isoprenoid quinones were extracted from the strain D13-10-4-6 T as reported by Collins et al. (1977), analyzed by high-performance liquid chromatography (Groth et al., 1997;Du et al., 2013), and confirmed by liquid chromatography/mass spectrometry. After culturing for 2 days in TSB at 30 • C, the cells were harvested at exponential phase and used for cellular fatty acids. Cellular fatty acids were extracted as reported by Kuykendall et al. (1988), analyzed using the Sherlock Microbial Identification System (Sasser, 1990).

Phenotypic Characterization
Growth conditions of the strain D13-10-4-6 T were determined at different temperature, pH, and salinity levels according to the method described by Li et al. (2016). The growth temperature was set at 4, 10,15,20,25,28,30,37,41, and 45 • C. The pH values for growth were adjusted to various pH values (pH 4.0-11.0, at intervals of 1.0 pH unit) by the buffers (Delory and King, 1945;Gomori, 1955) citrate and Na 2 HPO 4 buffer (pH 4.0-5.0), phosphate buffer (pH 6.0-7.0), Tris buffer (pH 8.0-9.0), and Na 2 HPO 4 /NaOH (pH 10.0-11.0). The salinity was determined in the range of 0-9% (w/v, intervals of 1%). Gram staining was performed according to the method described by Jenkins et al. (2003). To examine the anaerobic growth, the strain was incubated on TSA plates at 30 • C for 1 week in an anaerobic jar (Li et al., 2016). The activities of catalase and oxidase were determined by the methods described by Smibert and Krieg (1994). Enzymatic activity, carbon source utilization, and acid production were performed by API ZYM, API 20 NE, and API 50 CH (bioMérieux) according to the manufacturer's instructions, respectively.

Genome Information
The genome of strains D13-10-4-6 T and X. soli ZQBW T were sequenced and analyzed. In total, 63 contigs with a total sequence length of 4,683,906 bp for the strain X. soli ZQBW T were obtained, which was predicted to have 4,455 protein-coding genes, 47 tRNA genes, 3 rRNA genes, and 3 other RNA genes. The DNA G + C contents was 67.6%. The strain D13-10-4-6 T genome produced 66 contigs with a total sequence length of 4,605,234 bp, which was predicted to have 4,206 protein-coding genes, 45 tRNA genes, 3 rRNA genes, and 3 other RNA genes. The DNA G + C content of the strain D13-10-4-6 T was 62.9%, which was similar to P. bohemicus Cd-10 T (63.2%).

Phylogenetic Analyses
In this study, we have constructed phylogenetic trees based on the 16S rRNA gene, gyrB gene, and concatenated proteins (Figures 1-3) for representative members of the family Rhodobacteraceae encompassing 15 genera. The main groups clustering with the members of Rhodobacteraceae in 16S rRNA gene-based tree, gyrB gene-based tree, and concatenated proteins-based tree are almost consistent. The strains D13-10-4-6 T , P. bohemicus Cd-10 T , and X. humi IMT-291 T form one monophyletic group to in turn form Pseudogemmobacter clade with strong bootstrap support in all three phylogenetic trees (Figures 1-3), which is far removed from the branch of X. soli (the type species of the genus Xinfangfangia). X. humi IMT-291 T forms a distinct branch from the strains D13-10-4-6 T and P. bohemicus Cd-10 T in the Pseudogemmobacter clade. The results suggested that X. humi IMT-291 T should be a species belonging to the genus Pseudogemmobacter, although P. bohemicus Cd-10 T and X. humi IMT-291 T were published almost simultaneously and shared 99.26% 16S rRNA gene sequence similarity with each other. The strain D13-10-4-6 T forms a distinct branch from P. bohemicus Cd-10 T and X. humi IMT-291 T in all phylogenetic trees, and it has the highest 16S rRNA gene sequence similarity to P. bohemicus Cd-10 T (97.6%) and X. humi IMT-291 T (97.4%), and shares a less than 97% sequence similarity with all other validly published species. The results indicate that the strain D13-10-4-6 T should belong to a novel species of the genus Pseudogemmobacter.
Several genera are non-monophyletic, such as Rhodobacter, Tabrizicola, and Xinfangfangia. In most of the cases, the 16S rRNA gene-based tree shows its low discriminatory power. For instance, the species of the genus Tabrizicola are divided into two branches in the 16S rRNA gene-based tree, but they are clustered into three distinct branches, not least T. aquatica RCRI19 T (type species of the genus) and T. piscis K13M18 T are grouped together with R. thermarum YIM 73036 T , R. flagellatus SYSU G03088 T , and X. soli ZQBW T in trees based on gyrB gene and concatenated proteins tree with strong support. It can be seen from the trees based on the 16S rRNA gene, gyrB gene, and concatenated proteins that Rhodobacter and Tabrizicolaa are two closely related non-monophyletic genera. The members of the genus Tabrizicola are observed in three clades, clades A, B, and C, which are labeled in Figures 1-3. Clade A, formed by Tabrizicola alkalilacus DJC T , Tabrizicola sediminis DRYC-M-16 T , and Tabrizicola algicola ETT8 T , is next to the Gemmobacter clade and is far removed from two other Tabrizicola clades with a strong support in 16S rRNA genebased, gyrB gene-based, and concatenated proteins-based trees. These results suggest that clade A should belong to a novel genus of the family Rhodobacteraceae. Tabrizicola clade B, grouped by T. aquatica RCRI19 T (type species of the genus), T. piscis K13M18 T , R. thermarum YIM 73036 T , R. flagellatus SYSU G03088 T , and X. soli ZQBW T , is a monophyletic cluster found in trees based on the gyrB gene and concatenated proteins with a strong support, indicating that R. thermarum YIM 73036 T , R. flagellatus SYSU G03088 T , and X. soli ZQBW T should be transferred to the genus Tabrizicola. Tabrizicola fusiformis SY72 T , located in clade B in the 16S rRNA gene-based tree, is clustered together with Tabrizicola oligotrophica KMS-5 T to form clade C in both trees based on the gyrB gene and concatenated proteins, demonstrating that T. fusiformis SY72 T and T. oligotrophica KMS-5 T may be a novel genus of the family Rhodobacteraceae.
In the 16S rRNA gene-based tree, Cereibacter clade include R. alkalitolerans JA916 T , R. sediminicola JA983 T ), and members of the genus Cereibacter, except for Cereibacter changlensis JA139 T (the type species). The species C. changlensis JA139 T is observed in the Gemmobacter clade, which is similar to the results reported by Suresh et al. (2015). While C. changlensis JA139 T is grouped in the Cereibacter clade and located at the edge of the clade in trees inferred from the gyrB gene and concatenated proteins, indicating that it should belong to the genus Cereibacter, which is consistent to the results described by Suresh et al. (2015). R. sediminicola JA983 T is clustered in the Cereibacter clade in all the trees based on the 16S rRNA gene, gyrB gene, and concatenated proteins, indicating that they should be transferred to the genus Cereibacter.
The genus Rhodobacter proposed by Imhoff et al. (1984) contains 13 species with validated names according to the LSPN. In trees based on the 16S rRNA gene, gyrB gene, and concatenated protein, R. tardus CYK-10 T forms one distinct branch from other clades, suggesting that it should belong to a novel genus of the family Rhodobacteraceae. Three members (R. sediminicola JA983 T , R. thermarum YIM73036 T , R. flagellatus SYSU G03088 T ) are clustered into the Tabrizicola clade and Cereibacter clade, respectively. The other eight members of the genus Rhodobacter are temporarily classified into the genus Rhodobacter because of the absence of genome sequence of R. azollae, R. lacus, R. alkalitolerans, and R. sediminis, although they are grouped into two clusters in the 16S rRNA gene tree.

Genomic, Chemotaxonomic, and Physiological Analysis of the Novel Strain
The ANI values between the strain D13-10-4-6 T and its three closely related strains range from 74.4 to 81.2%, which are lower than the recommended ANI species boundary cutoff value (95-96%). The dDDH values between the strain D13-10-4-6 T and its closely related strains are 19.7-24.3%, lower than the threshold for species (70%). Those data indicate that the strain D13-10-4-6 T should belong to a novel species of the genus Pseudogemmobacter. Besides, P. bohemicus Cd-10 T and X. humi IMT-291 T share a 99.26% 16S rRNA gene sequence similarity, while their ANI and dDDH values are 79.1 and 22.1%, respectively (Table 1), which are lower than the species boundary cutoff values. Therefore, P. bohemicus Cd-10 T and X. humi IMT-291 T should belong to a different species of the genus Pseudogemmobacter.

Phylogenomic Metric Analysis
Average AAI is one of the well-established methods to separate prokaryotic genera (Luo et al., 2014;Rodriguez-R and Konstantinidis, 2014). It is proposed to be 65% AAI value for genera delineation of Bacteria and Archaea (Konstantinidis et al., 2017). However, the category thresholds of AAI for genus delineation are variable in many genera. For example, the value of 70% AAI is used to separate the genus Geomonas from the other genera of the family Geobacteraceae (Xu et al., 2020), and a range of 64.6-77.0% AAI is used to delineate different genera of the family Geobacteraceae (Xu et al., 2019). In this study, the values of AAI among 52 type strains from 15 related genera of the family Rhodobacteraceae have been determined and are listed in Table 4 and Supplementary Table 2. It can be seen from Table 4 that a gradient of 63.5-75.3% and 74.2-98.7% AAI values is found among the different clade (genera) and in the same clade of the family Rhodobacteraceae. Those data are consistent to the results of phylogeny based on concatenated proteins.  The Pseudogemmobacter clade, including strains D13-10-4-6 T , P. bohemicus Cd-10 T and X. humi IMT-291 T , has 77.4-81.8% AAI values among each other and shows 63.8-73.6% AAI values among the members of other clades in this study (Table 4 and Figure 4), which is consistent to the results of phylogeny based on the 16S rRNA gene, gyrB gene, and concatenated proteins (Figures 1-3). The Cereibacter clade, including R. sediminicola JA983 T and members of the genus Cereibacter, has 75.8-98.7% AAI values among the members of the clade and 65.2-72.7% AAI values among the members from the other clades in this work, indicating that R. sediminicola should be transferred to the genus Cereibacter (Table 4 and Supplementary Figure 1). Similarly, the AAI values within the Tabrizicola clade A and Tabrizicola clade B can also distinguish them from the other strains (Table 4 and Supplementary  Figures 2, 3).
Percentage of conserved protein is another method for genus delineation of prokaryote, and the value for genera delineation of POCP is proposed to be 50% (Qin et al., 2014). While the thresholds of POCP for genus delineation are also variable in many genera. The value of 65% POCP was used to separate the genus Geomonas from the other genus of the family Geobacteraceae (Xu et al., 2020), and most of the POCP values within the Roseobacter group comparisons were greater than 50% of the family Rhodobacteraceae (Wirth and Whitman, 2018). In this work, we also calculated the POCP values among the 52  (Table 4 and Figure 4). The same goes for the Cereibacter clade and Tabrizicola clade A (Table 4 and Supplementary Figures 1, 2).
The POCP values in this study are not always consistent to the phylogenetic analysis, as exemplified by the Tabrizicola (Supplementary Figure 3). The POCP values between X. soli ZQBW T (belonging to Tabrizicola clade B) and T. oligotrophica KMS-5 T , F. blasticum DSM 2131 T , and G. aestuarii CC-PW-75 T were slightly higher than those between X. soli ZQBW T and other members in Tabrizicola clade B (Supplementary Figure 3). Due to X. soli ZQBW T forming a stable clade within Tabrizicola clade B and forming a distinct clade with T. oligotrophica KMS-5 T , F. blasticum DSM 2131 T , and G. aestuarii CC-PW-75 T , we analyzed the pan-genome of Tabrizicola clade B in a supplementary analysis.
The pan-genome analysis was used in the classification of bacteria (Awan et al., 2018;Suresh et al., 2019). The amount of core genes was sensitive to heterogeneous in the coreand pan-genome analysis (Inglin et al., 2018;Suresh et al., 2019). The core gene numbers within were Tabrizicola clade B were considerably higher than those within the Tabrizicola clade B and T. oligotrophica KMS-5 T , F. blasticum DSM 2131 T , and G. aestuarii CC-PW-75 T (
The GenBank/EMBL/DDBJ accession numbers for the 16S rRNA gene and genome sequences for strain D13-10-4-6 T is MT036106 and JABJXT000000000, respectively.

Emended Description of the Genus Pseudogemmobacter
The description as given by Suman et al. (2019) remains correct except that the species are positive or negative for catalase and nitrate reductase.
Description of C. sediminicola Comb. nov.
The description of C. sediminicola is the same as that given for R. sediminicola by Suresh et al. (2020). The type strain is JA983 T (= KCTC 15782 T = NBRC 113843 T ).
Description of Stagnihabitans Gen. nov.
Cells are Gram-strain-negative, aerobic, non-motile, oxidasepositive, catalase-negative, ovoid to rod-shaped and divide by binary fission, sometimes forming chains. The predominant respiratory quinone is Q-10. The major cellular fatty acid is C 18:1 ω7c. PE, PG, and PC are the major polar lipids. The DNA G + C content is 66%. The member of the genus is separated from Rhodobacter based on the 16S rRNA, gyrB and concatenated protein phylogenetic trees, genome comparison. The type species is S. tardus comb. nov.
Description of S. tardus Comb. nov.
The description of S. tardus is the same as that given for R. tardus by Sheu et al. (2020). The type strain is CYK-10 T (= BCRC 81191 T = LMG 31336 T ).
Description of Pseudotabrizicola Gen. nov.
Cells are Gram-strain-negative, aerobic, non-motile, catalaseand oxidase-positive, rod-shaped. PG, DPG, PE, and PC are the major polar lipids. The predominant respiratory quinone is Q-10. The major cellular fatty acids are usually iso-C 18:0 , C 18:1 ω7c, and/or C 18:1 ω6c. The DNA G + C content is 62.9-64.4%. Members of the genus are separated from Tabrizicola based on the 16S rRNA, gyrB, and concatenated proteins phylogenetic trees, genome comparison. The type species is P. sediminis comb. nov.
Description of P. sediminis Comb. nov.
Description of P. algicola Comb. nov.
Description of T. thermarum Comb. nov.
Description of T. soli Comb. nov.
Tabrizicola soli (so'li. L. neut. n. soli of soil, the source of the type strain).
Basonym: Xinfangfangia soli (Hu et al., 2018). The description of T. soli is the same as that given for X. soli by Hu et al. (2018). The type strain is ZQBW T (= KCTC 62102 T = CCTCC AB 2017177 T ).

DATA AVAILABILITY STATEMENT
The datasets presented in this study can be found in online repositories. The names of the repository/repositories andaccession number(s) can be found below: NCBI-MT036106, JABJXT000000000, JAEACP000000000.

ACKNOWLEDGMENTS
We are grateful to Yan Liu, Fengjuan, Wang, and Wenjieng Li (China Forestry Culture Collection Center) for the material preparation and strain preservation.