Christiangramia qingdaonensis sp. nov., a novel polysaccharide-degrading Bacteroidota bacterium, isolated from intertidal sediment

Christiangramia qingdaonensis sp. nov., a novel polysaccharide-degrading Bacteroidota bacterium, isolated from intertidal sediment | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Christiangramia qingdaonensis sp. nov., a novel polysaccharide-degrading Bacteroidota bacterium, isolated from intertidal sediment Zhao Tong Chen, Bing Yu, Yu Shan Li, Zhi Wei Zhu, Xi Ying Zhang, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8043186/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 06 Jan, 2026 Read the published version in Antonie van Leeuwenhoek → Version 1 posted 11 You are reading this latest preprint version Abstract A novel polysaccharide-degrading bacterial strain, designated ASW11-125 T , was isolated from intertidal sediments in Aoshan Bay, Qingdao, China. The strain was strictly aerobic, Gram-stain-negative, catalase-positive but oxidase-negative, short rod-shaped, and exhibited gliding motility without flagella. Growth occurred at 4–35°C (optimum 28°C), pH 6.0–8.0 (optimum pH 7.0), and in 0.5–16.0% NaCl (optimum 2.5–3.0%). The predominant polar lipid was phosphatidylethanolamine. Major fatty acids were iso-C 15:0 and iso-C 17:0 3-OH, and the primary respiratory quinone was menaquinone-6 (MK-6). Based on the phylogenetic analyses of 16S rRNA gene sequences and 1542 single copy orthologous clusters, strain ASW11-125 T affiliated with the genus Christiangramia and was closely related to Christiangramia portivictoriae MCCC 1A00585 T (98.8%), Christiangramia aquimixticola KCTC 42706 T (98.8%) and Christiangramia marina KCTC 12366 T (98.6%). Comparative genomic analysis revealed that the average nucleotide identity (ANI) and digital DNA–DNA hybridization (dDDH) values between strain ASW11-125ᵀ and its closely related species (74.6–91.5% and 18.6–44.3%, respectively) were clearly lower than the proposed species cutoff values. Based on a polyphasic characterization integrating phenotypic, phylogenetic, and chemotaxonomic evidence, strain ASW11-125ᵀ represents a novel species of the genus Christiangramia , for which the name Christiangramia qingdaonensis sp. nov. is proposed. The draft genome of strain ASW11-125ᵀ is 3.2 Mb in size with a G + C content of 38.3%. Notably, genomic analysis revealed an abundance of genes encoding putative carbohydrate-active enzymes (CAZymes), particularly those associated with starch, laminarin, and fructan utilization, suggesting its potential role in the marine carbon cycle. The type strain is ASW11-125ᵀ (= KCTC 102340ᵀ = MCCC 1K09555ᵀ). Christiangramia qingdaonensis intertidal sediment polyphasic taxonomy polysaccharide utilization loci Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Marine phytoplankton fuel approximately 50% of Earth’s net primary production and exude a sizeable fraction as dissolved organic carbon (DOC) dominated by high-molecular-weight polysaccharides (Tang et al. 2017 ). Degradation of this reduced-carbon reservoir is disproportionately mediated by Bacteroidota , a lineage specialised in the enzymatic breakdown of structurally diverse marine polysaccharides (Lapébie et al. 2019 ; Zhu et al. 2023 ). From epipelagic waters to hadal trenches, marine Bacteroidota oscillate between three lifestyles: free-living, attached to algae cells (Mann et al. 2013 ), or embedded within sponges (Yoon and Oh 2012 ), corals (Sweet et al. 2011 ) and echinoderms (Romanenko et al. 2007 ; Gavriilidou et al. 2020 ). Gliding motility and rapid biofilm formation let them colonise different habitats (Mann et al. 2013 ). Once settled, polysaccharide-utilization loci (PULs) arm them with bespoke carbohydrate-active enzymes to unlock whatever polysaccharide the new niche offers. Each PUL forms a catabolic module of glycoside hydrolases, carbohydrate esterases, polysaccharide lyases and glycosyl-transferases coupled to TonB-dependent SusC transporters and SusD substrate-binding proteins, that synchronizes sensing, scission and import of oligosaccharides (Cantarel et al. 2009 ; Mann et al. 2013 ). Because the system is co-transcribed, cellular SusC/D abundance acts as an in-situ sentinel for which polysaccharide is being consumed at any given moment(Kappelmann et al. 2019 ). Even with the same PUL blueprint, sister Bacteroidota divide the glycan landscape at sub-genus resolution: free-living Hel1_33–49 keeps four PULs and a protease-heavy toolkit, while facultatively particle-attached Hel1_85 expands to eight PULs, swaps proteases for CAZymes, and rides living algae to mine a broader polysaccharide spectrum. This split in PUL repertoire epitomizes metabolic plasticity of Bacteroidota , reflecting their ability to adapt to diverse ecological niches (Xing et al. 2015 ). The genus Christiangramia , a member of the family Flavobacteriaceae , is a recognised taxon within the phylum Bacteroidota . Initially proposed by Nedashkovskaya et al. with Christiangramia echinicola as the type species (Nedashkovskaya et al. 2005 ), the genus currently comprises twenty-two validly described species, isolated from diverse marine habitats including tidal flats (Jeong et al. 2013 ; Park et al. 2015a ), seawater (Liu et al. 2014 ; Shahina et al. 2014 ; Shin et al. 2018 ), marine sediments (Hameed et al. 2014 ; Yoon et al. 2015 ; Li et al. 2018 ; Yang et al. 2023 ), solar salterns (Joung et al. 2011 ), and marine organisms (Nedashkovskaya et al. 2005 ; Nedashkovskaya et al. 2010 ; Liu et al. 2020 ). Most members are Gram-stain-negative, rod-shaped, strictly aerobic or aerobic, exhibit gliding motility, and contain menaquinone-6 (MK-6) as the major respiratory quinone (Yang et al. 2023 ; Wang et al. 2025 ). The genomic DNA G + C content ranges from 36.6 to 48.0% (Park et al. 2015b ; Panschin et al. 2017 ). Functionally, Christiangramia species arm each niche with bespoke CAZyme batteries. C. forsetii KT0803 packs 164 CAZymes into starch-, laminarin- and alginate-specific PULs (Kabisch et al. 2014 ), whereas C. flava JLT2011 swaps alginate for two xylan PULs and a pectin PUL, parking half of its GH genes inside these loci (Tang et al. 2017 ). Multi-omics show that such differences in PUL composition directly influence substrate utilization patterns. Yet we still lack a genus-wide map of which loci are universal, which are niche-exclusive, and how this mosaic quantitatively gates Christiangramia ’s share of marine carbon flux. During an investigation of bacterial diversity in the Qingdao intertidal zone, a novel strain, ASW11-125ᵀ, was isolated from the sediment and assigned to the genus Christiangramia . To determine its precise taxonomic status and explore its functional potential, a polyphasic taxonomic characterization was conducted, including physiological, chemotaxonomic, and phylogenetic analyses, as well as genomic analyses. The results confirmed that strain ASW11-125ᵀ represents a novel species equipped with a private suite of starch-, laminarin- and fructan-specific PULs. Within the genus, laminarin and starch are universal metabolic pillars; fructan utilization is an accessory module, switched on only by those lineages that encounter it. Materials and methods Sample collection and bacterial isolation A sediment sample was collected from the tidal flat of Aoshan Bay (36°22′1″N, 120°41′51″E) in Qingdao, China. Approximately 1 g of sediment was serially diluted in 3% sterile artificial seawater (Sigma, USA), and appropriate dilutions were spread onto 1/3-strength marine agar 2216 (MA). After incubation at 20°C for 7 days, a single yellow colony was picked and purified through repeated subculturing on fresh plates, designated ASW11-125ᵀ. For long-term storage, the strain was cultured in marine broth 2216 (MB) at 28°C and preserved at − 80°C in MB supplemented with 20% (v/v) glycerol. For further comparative taxonomic analysis, the reference strains C. portivictoriae MCCC 1A00585 T , C. aquimixticola KCTC 42706 T and C. marina KCTC 12366 T were included in all subsequent experiments under the same conditions. Phylogenetic analysis The 16S rRNA gene of strain ASW11-125 T was amplified via PCR with the universal primers 27F (5′-AGAGTTTGATCCTGGCTCAG-3′) and 1492R (5′-GGYTACCTTGTTACGACTT-3′). The resulting nearly full-length sequence (1491 bp) was compared with related strains using the EzBioCloud database and the BLAST algorithm. Multiple sequence alignments were performed with the CLUSTAL W algorithm (Thompson et al. 1994 ) in MEGA 11 (Tamura et al. 2021 ). Phylogenetic trees were constructed using the maximum-likelihood (Felsenstein 1981 ), neighbor-joining (Saitou and Nei 1987 ), and minimum-evolution methods (Pardi et al. 2010 ) in MEGA 11, with evolutionary distances calculated under the Kimura two-parameter model (Kimura 1980 ). Bootstrap analyses were performed based on 1000 replicates to evaluate the robustness of tree topologies (Felsenstein 1985 ). For the genome-based phylogenetic analysis, genomic sequences of closely related species were obtained from GenBank. The 1542 single-copy orthologous clusters (OCs) from 25 bacterial genomes were selected with Proteinortho v5.16 (Lechner et al. 2011 ) and aligned with MUSCLE v3.8.31 (Edgar 2004 ). Phylogenetic trees were subsequently inferred from the concatenated alignment of these clusters using MEGA 11. Physiological and biochemical analysis Colony morphology of strain ASW11-125 T was observed on MA after 48 h of incubation at 28°C. For detailed morphology features, including cell size, shape and flagella, transmission electron microscopy was used with cells negatively stained with 3% uranyl acetate for 1 min. Gram staining was performed using a Gram-stain kit (Solarbio, China) following the manufacturer’s instructions. Gliding motility was examined as described by Bowman ( 2000 ) and flexirubin-type pigment production was evaluated according to Bernardet and Bowman ( 2006 ). Growth under different pH conditions was assessed in marine broth (MB) adjusted to pH 4.0–10.0 (in increments of 0.5 units) and pH 10.0–12.0 (in increments of 1.0 unit) using the following buffers (50 mM each): MES (pH 4.0–6.0), MOPS (pH 6.5–7.0), Tris (pH 7.5–8.5), CHES (pH 9.0–10.0), and CAPS (pH 11.0–12.0). NaCl tolerance was assessed in NaCl-free basal MB supplemented with 0–20% NaCl (0–4% at 0.5% intervals; 4–20% at 1% intervals). The temperature range for growth was examined in MB at 4, 15, 20, 25, 28, 32, 35, 37 and 40°C, respectively. Anaerobic growth was evaluated on modified TYS broth (0.5% tryptone, 0.1% yeast extract, 3.0% sea salt) supplemented with 0.15% (w/v) cysteine hydrochloride and 1‰ (w/v) resazurin for 1 month in an anaerobic jar. Catalase activity was determined by bubble production in 3% (v/v) H 2 O 2 , and oxidase activity was assessed using commercial test strips (Merck). Hydrolysis of milk (0.5%, w/v), casein (0.5%, w/v), gelatin (1%, w/v), starch (1%, w/v), cellulose (1%, w/v), algin (1%, w/v) and Tweens 20, 40, 60, and 80 (1%, v/v) were carried out as described by Smibert and Krieg ( 1981 ). Antibiotic susceptibility was tested by the disc diffusion method (Hudzicki 2009 ) on MA plates using the following antibiotics: ampicillin (10 µg), cephalexin (30 µg), erythromycin (15 µg), roxithromycin (15 µg), oxytetracycline (30 µg), streptomycin (10 µg), vancomycin (30 µg), polymyxin B (300 IU), sulfadiazine (25 µg), sulfamethoxazole (25 µg), sulfafurazole (25 µg), norfloxacin (10 µg), levofloxacin (5 µg), lincomycin (2 µg) and chloramphenicol (30 µg). Additional enzyme activities and biochemical properties were examined by API ZYM and API 20NE test strips (bioMérieux), following standard protocols with a two-week extended incubation. Bacterial suspensions were prepared in 3% NaCl for API ZYM and in AUX medium for API 20NE, respectively. Chemotaxonomic analysis For cellular fatty acid analysis, the cells of strain ASW11-125 T and the reference strains were cultured in marine broth (MB) at 28°C and harvested during the early exponential growth phase (12–24 h). Fatty acid methyl esters were prepared from lyophilized cells by saponification, methylation, and extraction following the standard protocol of the Sherlock Microbial Identification System (MIDI). They were then analysed using the Microbial Identification System (MIS) software for identification and quantification. Polar lipids and isoprenoid quinones were extracted from cells collected at the late exponential phase under the same culture conditions. Polar lipids were extracted according to the method of Komagata and Suzuki ( 1988 ) and separated by two-dimensional thin-layer chromatography. Specific lipids were detected by spraying with the following reagents: phosphomolybdic acid (for total lipids), ninhydrin (for aminolipids), molybdenum blue (for phospholipids) (Collins and Jones 1980 ). Respiratory quinones were analysed from lyophilized cells using a chloroform/methanol (2:1, v/v) extraction system. The extract was examined using a Dionex Ultimate 3000 HPLC system coupled to a Bruker impact HD mass spectrometer for identification. Genomic sequencing and analysis The genome of strain ASW11-125 T was sequenced on an Illumina HiSeq X-ten platform (OE Biotech, Shanghai, PR China). Raw reads were quality-controlled using Trimmomatic and assembled using ABySS version 2.0. The assembled draft genome was submitted to the GenBank database. The DNA G + C content was calculated from the genome sequence. Genome relatedness was assessed by the average nucleotide identity (ANI) using the EzBioCloud tool ( https://www.ezbiocloud.net/tools/ani ) (Yoon et al. 2017 ), and digital DNA-DNA hybridization (dDDH), estimated through the GGDC web server ( http://ggdc.dsmz.de/ggdc.php ) (Meier-Kolthoff et al. 2013 ).Functional annotation was performed using the RAST server (Aziz et al. 2008 ). To assess the polysaccharide-degrading potential, carbohydrate-active enzymes (CAZymes) were identified using the HMMER tool in the dbCAN3 server (Zheng et al. 2023 ). Polysaccharide-utilization loci (PULs) were further predicted by searching the dbCAN-PUL database. Variation in glycoside hydrolase (GH) composition across different genera was visualised using non-metric multidimensional scaling (NMDS) based on Bray-Curtis dissimilarities. Results and discussion Phylogenetic analysis The nearly complete 16S rRNA gene sequence of strain ASW11-125 T (1491 bp; GenBank accession no. PV134465) was obtained in this study. Comparative analysis revealed that strain ASW11-125ᵀ belongs to the genus Christiangramia , showing the highest sequence similarity to C. portivictoriae MCCC 1A00585 T (98.8%), followed by C. aquimixticola KCTC 42706 T (98.8%), C. marina KCTC 12366 T (98.6%), and other validly published type strains of this genus (93.3%–98.3%). In the ML, NJ and ME phylogenetic trees (Figs. 1 , S1, and S2), strain ASW11-125 T consistently formed a robust, distinct clade together with C. portivictoriae MCCC 1A00585 T , C. aquimixticola KCTC 42706 T and C. marina KCTC 12366 T . This subclade was stably positioned within a larger clade comprising all recognised species of the genus Christiangramia , with strong bootstrap support (> 89%). Phylogenomic trees based on 1542 single-copy OCs confirmed that strain ASW11-125ᵀ, C. marina KCTC 12366 T and C. portivictoriae MCCC 1A00585 T formed a subclade, which further clustered with C. aquimixticola KCTC 42706 T into a larger clade (Figs. 2 , S3, and S4). Genomic relatedness analysis indicated that the average nucleotide identity (ANI) and digital DNA–DNA hybridization (dDDH) values between strain ASW11-125ᵀ and the closely related strains were 91.5%, 85.2%, and 74.6% (ANI), and 44.3%, 28.7%, and 18.6% (dDDH), respectively (Table S1 ). All values fell well below the accepted species demarcation thresholds (ANI < 95–96%; dDDH < 70%), supporting the recognition of strain ASW11-125ᵀ as a novel species (Meier-Kolthoff et al. 2013 ; Palmer et al. 2020 ). Phenotypic and biochemical characterization Strain ASW11-125ᵀ was determined to be Gram-stain-negative, strictly aerobic, and capable of gliding motility. Flexirubin-type pigments were not produced. Transmission electron microscopy showed that the cells were short rods, approximately 0.5–0.8 µm wide and 1.0–3.0 µm long, and lacked flagella (Fig. S5). After 48 h of incubation on MA at 28°C, colonies were round, smooth, yellow, and 0.5–1.0 mm in diameter. The strain grew at 4–35°C (optimum 28°C), pH 6.0–8.0 (optimum pH 7.0) and in the presence of 0.5–16.0% NaCl (optimum 2.5–3.0%). In antibiotic susceptibility tests, the isolate was resistant to ampicillin, cephalexin, erythromycin, streptomycin, sulfadiazine, and sulfafurazole, but susceptible to roxithromycin, oxytetracycline, sulfamethoxazole, norfloxacin, levofloxacin, vancomycin, polymyxin B, lincomycin, and chloramphenicol. A detailed comparison of the physiological and biochemical characteristics of strain ASW11-125ᵀ and its phylogenetic relatives is provided in the species description and in Table 1 . Table 1 Differential characteristics of strains ASW11-125 T and type strains of closely related species Characteristic 1 2 3 4 5 6 Growth range of: Temperature (optimum, ℃) 4–35 (28) 4–37 (28–30) † 4–36 (28–30) † 10–40 (30) † 4–37 (23–25) 4–40 (30) NaCl (optimum, %) 0.5–16 (2.5–3) 1–15 (2–5) † 1–6 † 0.5–8 (1–2) † 1–15 (4–5) * 0–9 (2) pH (optimum) 6.0–8.0 (7.0) ND † 6.0–10.0 (7.0–8.0) † 6.0–ND (7.0–8.0) † 5–9 (7–7.5) * 6–ND (7.0–8.0) Acid production from: Glucose – + – – + ND Utilization of: Arginine – – – – ND ND Arabinose – + + – + – Mannose – – – – – – Maltose – + – – + – Malate + + – – ND – Urea – – – – – – Hydrolysis of: Tween20 + + + – – ND Tween40 + + + – + ND Tween80 W + + – + + Starch + + + – + + Enzyme activity (API ZYM): Lipase (C14) + + + + + * – Trypsin + + + + w * – β -Galactosidase – + – – + – α -Glucosidase – + + + + * – N-acetyl glucosaminidase – + – – + * + DNA G + C content (%) 38.3 40.0 † 39.9 † 48.0 † 39.6 39.2 Strains: 1, C. qingdaonensis ASW11-125 T ; 2, C. marina KCTC 12366 T ; 3, C. portivictoriae MCCC 1A00585 T ; 4, C. aquimixticola KCTC 42706 T ; 5, C. echinicola KMM 6050 T ; 6, C. sabulilitoris HSMS-1 T . All strains were Gram-negative and tested positive for catalase activities; hydrolysis of gelatin; and enzyme activities of alkaline phosphatase, esterase (C4), esterase lipase (C8), leucine arylamidase, valine arylamidase, cystine arylamidase, acid phosphatase and naphthol-AS-BI-phosphohydrolase. All strains were negative for citrate utilization. Data for columns 1–4 were obtained in this study, while data for columns 5–6 were taken from Nedashkovskaya et al. ( 2005 ), Park et al. ( 2020 ), respectively. +, positive; –, negative; ND, no data available; W, weakly positive; *, Data from Li et al. 2018 ; †, Data from Lau et al. 2005 , Nedashkovskaya et al. 2010 , Park et al. 2015b . Chemotaxonomic characteristics The major fatty acids (≥ 10%) of strain ASW11-125ᵀ were iso-C 15:0 (22.7%) and iso-C 17:0 3-OH (10.0%), which are consistent with the general profile of the genus Christiangramia . However, notable differences were observed in the proportions of several fatty acids compared to closely related strains. For example, strain ASW11-125ᵀ exhibited significantly lower proportion of C 16:0 (5.5%) than C. portivictoriae MCCC 1A00585ᵀ (11.5%). Similarly, its content of anteiso-C 15:0 (7.5%) was substantially lower than that in C. aquimixticola KCTC 42706ᵀ (14.8%) (Table 2 ). Polar lipid analysis indicated that strain ASW11-125 T , along with C. portivictoriae MCCC 1A00585 T , C. aquimixticola KCTC 42706 T and C. marina KCTC 12366 T , all included phosphatidylethanolamine (PE) and two unidentified lipids (L1, L2). An additional unidentified lipid (L3) was detected in both ASW11-125ᵀ and C. portivictoriae MCCC 1A00585ᵀ, while an aminolipid (AL) was present only in C. marina KCTC 12366ᵀ (Fig. S6). The respiratory quinone of strain ASW11-125 T was MK-6, which aligns with the typical quinone system of the genus Christiangramia . Table 2 Cellular fatty acid compositions (%) of strain ASW11-125 T and closely related strains of the genus Christiangramia Fatty acid 1 2 3 4 Straight-chain C 16:0 5.5 9.3 11.5 6.0 C 18:0 2.7 3.6 6.0 2.8 Branched iso-C 14:0 1.3 1.0 ND ND iso-C 15:0 22.7 21.8 19.2 14.8 iso-C 15:1 1.2 1.5 2.4 0.8 iso-C 16:0 9.2 6.8 4.1 4.9 iso-C 16:1 H 3.2 1.5 1.6 2.0 anteiso-C 15:0 7.5 7.8 8.2 14.8 Hydroxy C 15:0 2-OH 2.1 2.3 1.9 2.1 C 17:0 2-OH 3.2 3.7 3.5 7.3 iso-C 15:0 3-OH 1.9 1.7 1.8 1.0 iso-C 16:0 3-OH 4.0 2.4 2.2 1.7 iso-C 17:0 3-OH 10.0 11.7 10.6 7.8 Unsaturated C 15:1 ω 6 c 1.5 0.5 1.5 0.8 C 17:1 ω 6 c 2.8 1.2 1.2 1.8 C 17:1 ω 8 c 0.8 1.0 0.5 ND anteiso-C 17:1 ω 9 c 2.0 1.1 1.9 8.3 Summed features* 3 8.9 9.7 8.6 9.3 9 5.4 6.0 5.2 8.7 Strains: 1, C. qingdaonensis ASW11-125 T ; 2, C. marina KCTC 12366 T ; 3, C. portivictoriae MCCC 1A00585 T ; 4, C. aquimixticola KCTC 42706 T . All data were obtained from this study. Major fatty acids in each strain are shown in bold. *Summed features are fatty acids that cannot be resolved reliably from another fatty acid using the chromatographic conditions chosen. The MIDI system reports these as a combined feature with a single percentage of the total; Summed feature 3 contains C 16:1 ω 7 c and/or C 16:1 ω 6 c and/or iso-C 15:0 2-OH; Summed feature 9 contains iso-C 17:1 ω 9 c and/or C 16:0 10-methyl. ND, no data available. Genome features The draft genome of strain ASW11-125 T consists of 3,203,245 bp with 19 contigs, an N50 value of 880,254 bp, and a DNA G + C content of 38.3%. A total of 3,010 genes were predicted, including 2981 protein-coding genes, 6 pseudogenes, and 23 RNA genes (3 rRNA genes, 16 tRNA genes, and 4 other RNA genes) (Table S2). Genomic comparisons indicated that strain ASW11-125ᵀ had significantly higher gene abundances for amino acid/derivative metabolism (21.4%) and cofactor/vitamin/pigment metabolism (15.2%), but maintained comparable levels for protein (11.5%), carbohydrate (9.1%), and nucleoside/nucleotide (7.2%) metabolism relative to its close relatives (Table S3). Based on dbCAN3 analysis, 131 carbohydrate-active enzymes (CAZymes) were identified, accounting for 4.35% of the protein-coding sequences (Table S4). These included 68 glycosyltransferases (GTs, 16 families), 40 glycoside hydrolases (GHs, 25 families), 14 carbohydrate esterases (CEs, 4 families), 2 auxiliary activities (AAs, 2 families), and 7 carbohydrate-binding modules (CBMs, 6 families). NMDS analysis based on GH profiles revealed that the genus Christiangramia clustered within a group of known polysaccharide-degrading Bacteroidota , including Polaribacter , Flavobacterium , Bacteroides , and Algibacter , and was clearly separated from the non-saccharolytic genus Staphylococcus (ANOSIM, r = 0.66, P = 0.001; Fig. 3 a). Furthermore, Christiangramia exhibited significantly greater similarity to Algibacter and Polaribacter than to other genera ( p < 0.05) (Fig. 3 b), indicating a closely aligned potential for polysaccharide degradation. Polysaccharide-utilization loci for starch, laminarin and fructan Strain ASW11-125ᵀ harbors three polysaccharide utilization loci (PULs) that target starch, laminarin, and fructan. Each locus encodes a canonical PUL architecture, consisting of CAZymes, SusC/D-like transporters, and transcriptional regulators—an arrangement that is conserved across Christiangramia species (Fig. 4 ). Synteny across 18 Christiangramia genomes showed the laminarin locus in every strain and starch loci in 94%, whereas the fructan locus is confined to one-third (33%) (Fig. 4 ). This distribution suggests that laminarin and starch are genus-wide metabolic pillars; fructan utilization constitutes a facultative trait likely maintained in specific ecological contexts. Collectively, these PULs deliver a rapid high-molecular-weight carbon uptake module that confers ecological plasticity and competitive supremacy throughout marine ecosystems. Taxonomic conclusion The phenotypic, phylogenetic, biochemical, genomic and chemotaxonomic features all support that strain ASW11-125 T represents a novel species of the genus Christiangramia , for which the name Christiangramia qingdaonensis sp. nov. is proposed here. Description of Christiangramia qingdaonensis sp. nov. Christiangramia qingdaonensis [qing.dao.nen’sis. N.L. fem. adj. qingdaonensis, pertaining to Qingdao, China, where the type strain was isolated] Cells are strictly aerobic, Gram-stain-negative, non-spore-forming, rod-shaped, non-flagellated and motile by gliding, measuring 0.5–0.8 µm in width and 1.0–3.0 µm in length. After 48 h on marine agar at 28°C, colonies are yellow, smooth, and 0.5–1.0 mm in diameter. Growth occurs at 4–35°C (optimally at 28°C), pH 6.0–8.0 (optimally at pH 7.0) and in the presence of 0.5–16.0% (w/v) NaCl (optimum 2.5–3.0%). Flexirubin-type pigments are not produced. Cells are catalase-positive but oxidase-negative. Hydrolytic activities are observed for Tweens 20, 40, 60, and 80, starch, aesculin, casein, and gelatin, but not for cellulose or milk. Nitro- β -D-methylgalactose and malate are assimilated, while arabinose, mannose, mannitol, maltose, and citrate are not. Enzymatic activities are positive for alkaline phosphatase, esterase (C4), esterase lipase (C8), lipase (C14), leucine arylamidase, valine arylamidase, cysteine arylamidase, trypsin, chymotrypsin, acid phosphatase, naphthol-AS-BI-phosphohydrolase, α -galactosidase, and β -glucosidase. However, activities are negative for urease, arginine dihydrolase, β -galactosidase, α -glucosidase, and β -fucosidase. Acid production from glucose, nitrate reduction, and indole production are negative. The major polar lipid is phosphatidylethanolamine, and the predominant fatty acids are iso-C 15:0 (22.7%) and iso-C 17:0 3-OH (10.0%). The respiratory quinone is menaquinone-6 (MK-6). The genomic DNA G + C content of the type strain is 38.3%. The type strain, ASW11-125 T (= KCTC 102340 T = MCCC 1K09555 T ), was isolated from intertidal sediment of Qingdao, PR China. The GenBank accession numbers for the 16S rRNA gene and whole-genome sequence are PV134465 and NZ_JBLWIM000000000, respectively. Abbreviations MCCC Marine culture collection of China KCTC Korean collection for type cultures MEGA Molecular evolutionary genetics analysis RAST Rapid annotations using subsystems technology ANI Average nucleotide identity dDDH Digital DNA-DNA hybridization Declarations Competing interests The authors declare no competing interests. Ethics approval This article does not contain any studies conducted by the authors involving human participants or animals. Funding This work was funded by the National Natural Science Foundation of China (32300115). Author Contribution ZTC designed the study, performed the data analysis, and wrote the manuscript. BY, YSL and ZWZ conducted genome analysis, contributed to the interpretation of data, and revised the manuscript. YL and XYZ supervised the experimental work, provided feedback on the manuscript, and contributed to the final manuscript revision. 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14:01:10","extension":"html","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":162782,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/98b460038189780a44c8c1d6.html"},{"id":96918727,"identity":"cd68ac3f-24b0-47a8-9f12-2cf220a921c1","added_by":"auto","created_at":"2025-11-27 14:12:26","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":268504,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum-likelihood phylogenetic tree based on 16S rRNA gene sequences, showing the phylogenetic positions of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e (in bold) and the phylogenetically related species. Bootstrap values (\u0026gt; 50%) based on 1000 replicates are shown at nodes. Filled circles indicate branches that were also recovered in both the neighbor-joining tree and the minimum-evolution tree. Bar, 0.02 substitutions per nucleotide position\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/f081ff7bceb78eb923d42335.png"},{"id":96831894,"identity":"c1ac4230-c5c5-4ae0-896a-5db47f4c8816","added_by":"auto","created_at":"2025-11-26 14:01:09","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":240444,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum-likelihood phylogenomic tree based on 1542 single-copy orthologous cluster sequences from genome sequences showing phylogenetic positions of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e and related species in the family \u003cem\u003eFlavobacteriaceae\u003c/em\u003e. Filled circles indicate branches that were also recovered in both the neighbor­joining tree and the minimum-evolution tree. Bar, 0.05 substitutions per amino acid position\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/54d5ddbff1f2ae7cafd46f02.png"},{"id":96918485,"identity":"b1c83a2c-7a0e-48f8-8cb2-9f0722b06710","added_by":"auto","created_at":"2025-11-27 14:12:00","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":246224,"visible":true,"origin":"","legend":"\u003cp\u003eVariation in glycoside hydrolase (GH) composition across different genera. (a) NMDS ordination of GH profiles based on Bray-Curtis dissimilarities. ANOSIM results: R = 0.662, P = 0.001. (b) Bray-Curtis dissimilarity comparisons between different genera: 1. \u003cem\u003eStaphylococcus\u003c/em\u003e (\u003cem\u003eSta\u003c/em\u003e); 2. \u003cem\u003eFlavobacterium\u003c/em\u003e (\u003cem\u003eFla\u003c/em\u003e); 3. \u003cem\u003ePolaribacter\u003c/em\u003e (\u003cem\u003ePol\u003c/em\u003e); 4. \u003cem\u003eBacteroides\u003c/em\u003e (\u003cem\u003eBac\u003c/em\u003e); 5. \u003cem\u003eAlgibacterc\u003c/em\u003e (\u003cem\u003eAlg\u003c/em\u003e); 6. \u003cem\u003eChristiangramia\u003c/em\u003e(\u003cem\u003eChr\u003c/em\u003e). The different letters (i.e. “a”, “b”, “c” and “d” in box plots) represent significant differences (\u003cem\u003ep\u003c/em\u003e \u0026lt; 0.05) based on one-way ANOVA followed by Tukey's HSD test\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/4648bf71b95483f90a0467ee.png"},{"id":96918667,"identity":"07d0c493-4670-4a6e-855e-02cd26b6f2dc","added_by":"auto","created_at":"2025-11-27 14:12:17","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":535185,"visible":true,"origin":"","legend":"\u003cp\u003eDistribution of starch, laminarin, and fructan utilization loci (PULs) across \u003cem\u003eChristiangramia\u003c/em\u003e species\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/ae5513d54bb000789f258533.png"},{"id":100070278,"identity":"3b3165ab-94af-45dc-8c7f-70dd2737664b","added_by":"auto","created_at":"2026-01-12 16:17:21","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2973719,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/ba837251-189c-497e-b458-0d81694ad782.pdf"},{"id":96831899,"identity":"3eef81f9-89de-4cfc-9c29-1a3b88570ede","added_by":"auto","created_at":"2025-11-26 14:01:09","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":3067125,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarymaterials.docx","url":"https://assets-eu.researchsquare.com/files/rs-8043186/v1/ce819fd5dfb1db4207a4b69f.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Christiangramia qingdaonensis sp. nov., a novel polysaccharide-degrading Bacteroidota bacterium, isolated from intertidal sediment","fulltext":[{"header":"Introduction","content":"\u003cp\u003eMarine phytoplankton fuel approximately 50% of Earth\u0026rsquo;s net primary production and exude a sizeable fraction as dissolved organic carbon (DOC) dominated by high-molecular-weight polysaccharides (Tang et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Degradation of this reduced-carbon reservoir is disproportionately mediated by \u003cem\u003eBacteroidota\u003c/em\u003e, a lineage specialised in the enzymatic breakdown of structurally diverse marine polysaccharides (Lap\u0026eacute;bie et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Zhu et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). From epipelagic waters to hadal trenches, marine \u003cem\u003eBacteroidota\u003c/em\u003e oscillate between three lifestyles: free-living, attached to algae cells (Mann et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), or embedded within sponges (Yoon and Oh \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), corals (Sweet et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and echinoderms (Romanenko et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Gavriilidou et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Gliding motility and rapid biofilm formation let them colonise different habitats (Mann et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Once settled, polysaccharide-utilization loci (PULs) arm them with bespoke carbohydrate-active enzymes to unlock whatever polysaccharide the new niche offers. Each PUL forms a catabolic module of glycoside hydrolases, carbohydrate esterases, polysaccharide lyases and glycosyl-transferases coupled to TonB-dependent SusC transporters and SusD substrate-binding proteins, that synchronizes sensing, scission and import of oligosaccharides (Cantarel et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Mann et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Because the system is co-transcribed, cellular SusC/D abundance acts as an in-situ sentinel for which polysaccharide is being consumed at any given moment(Kappelmann et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Even with the same PUL blueprint, sister \u003cem\u003eBacteroidota\u003c/em\u003e divide the glycan landscape at sub-genus resolution: free-living Hel1_33\u0026ndash;49 keeps four PULs and a protease-heavy toolkit, while facultatively particle-attached Hel1_85 expands to eight PULs, swaps proteases for CAZymes, and rides living algae to mine a broader polysaccharide spectrum. This split in PUL repertoire epitomizes metabolic plasticity of \u003cem\u003eBacteroidota\u003c/em\u003e, reflecting their ability to adapt to diverse ecological niches (Xing et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe genus \u003cem\u003eChristiangramia\u003c/em\u003e, a member of the family \u003cem\u003eFlavobacteriaceae\u003c/em\u003e, is a recognised taxon within the phylum \u003cem\u003eBacteroidota\u003c/em\u003e. Initially proposed by Nedashkovskaya et al. with \u003cem\u003eChristiangramia echinicola\u003c/em\u003e as the type species (Nedashkovskaya et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), the genus currently comprises twenty-two validly described species, isolated from diverse marine habitats including tidal flats (Jeong et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Park et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2015a\u003c/span\u003e), seawater (Liu et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Shahina et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Shin et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), marine sediments (Hameed et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Yoon et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Yang et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), solar salterns (Joung et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and marine organisms (Nedashkovskaya et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Nedashkovskaya et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Most members are Gram-stain-negative, rod-shaped, strictly aerobic or aerobic, exhibit gliding motility, and contain menaquinone-6 (MK-6) as the major respiratory quinone (Yang et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The genomic DNA G\u0026thinsp;+\u0026thinsp;C content ranges from 36.6 to 48.0% (Park et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015b\u003c/span\u003e; Panschin et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Functionally, \u003cem\u003eChristiangramia\u003c/em\u003e species arm each niche with bespoke CAZyme batteries. \u003cem\u003eC. forsetii\u003c/em\u003e KT0803 packs 164 CAZymes into starch-, laminarin- and alginate-specific PULs (Kabisch et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), whereas \u003cem\u003eC. flava\u003c/em\u003e JLT2011 swaps alginate for two xylan PULs and a pectin PUL, parking half of its GH genes inside these loci (Tang et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Multi-omics show that such differences in PUL composition directly influence substrate utilization patterns. Yet we still lack a genus-wide map of which loci are universal, which are niche-exclusive, and how this mosaic quantitatively gates \u003cem\u003eChristiangramia\u003c/em\u003e\u0026rsquo;s share of marine carbon flux.\u003c/p\u003e\u003cp\u003eDuring an investigation of bacterial diversity in the Qingdao intertidal zone, a novel strain, ASW11-125ᵀ, was isolated from the sediment and assigned to the genus \u003cem\u003eChristiangramia\u003c/em\u003e. To determine its precise taxonomic status and explore its functional potential, a polyphasic taxonomic characterization was conducted, including physiological, chemotaxonomic, and phylogenetic analyses, as well as genomic analyses. The results confirmed that strain ASW11-125ᵀ represents a novel species equipped with a private suite of starch-, laminarin- and fructan-specific PULs. Within the genus, laminarin and starch are universal metabolic pillars; fructan utilization is an accessory module, switched on only by those lineages that encounter it.\u003c/p\u003e"},{"header":"Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003eSample collection and bacterial isolation\u003c/h2\u003e\u003cp\u003eA sediment sample was collected from the tidal flat of Aoshan Bay (36\u0026deg;22\u0026prime;1\u0026Prime;N, 120\u0026deg;41\u0026prime;51\u0026Prime;E) in Qingdao, China. Approximately 1 g of sediment was serially diluted in 3% sterile artificial seawater (Sigma, USA), and appropriate dilutions were spread onto 1/3-strength marine agar 2216 (MA). After incubation at 20\u0026deg;C for 7 days, a single yellow colony was picked and purified through repeated subculturing on fresh plates, designated ASW11-125ᵀ. For long-term storage, the strain was cultured in marine broth 2216 (MB) at 28\u0026deg;C and preserved at \u0026minus;\u0026thinsp;80\u0026deg;C in MB supplemented with 20% (v/v) glycerol. For further comparative taxonomic analysis, the reference strains \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e, \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e and \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e were included in all subsequent experiments under the same conditions.\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003ePhylogenetic analysis\u003c/h3\u003e\n\u003cp\u003eThe 16S rRNA gene of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e was amplified via PCR with the universal primers 27F (5\u0026prime;-AGAGTTTGATCCTGGCTCAG-3\u0026prime;) and 1492R (5\u0026prime;-GGYTACCTTGTTACGACTT-3\u0026prime;). The resulting nearly full-length sequence (1491 bp) was compared with related strains using the EzBioCloud database and the BLAST algorithm. Multiple sequence alignments were performed with the CLUSTAL W algorithm (Thompson et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e1994\u003c/span\u003e) in MEGA 11 (Tamura et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Phylogenetic trees were constructed using the maximum-likelihood (Felsenstein \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1981\u003c/span\u003e), neighbor-joining (Saitou and Nei \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1987\u003c/span\u003e), and minimum-evolution methods (Pardi et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) in MEGA 11, with evolutionary distances calculated under the Kimura two-parameter model (Kimura \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). Bootstrap analyses were performed based on 1000 replicates to evaluate the robustness of tree topologies (Felsenstein \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). For the genome-based phylogenetic analysis, genomic sequences of closely related species were obtained from GenBank. The 1542 single-copy orthologous clusters (OCs) from 25 bacterial genomes were selected with Proteinortho v5.16 (Lechner et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) and aligned with MUSCLE v3.8.31 (Edgar \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Phylogenetic trees were subsequently inferred from the concatenated alignment of these clusters using MEGA 11.\u003c/p\u003e\n\u003ch3\u003ePhysiological and biochemical analysis\u003c/h3\u003e\n\u003cp\u003eColony morphology of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e was observed on MA after 48 h of incubation at 28\u0026deg;C. For detailed morphology features, including cell size, shape and flagella, transmission electron microscopy was used with cells negatively stained with 3% uranyl acetate for 1 min. Gram staining was performed using a Gram-stain kit (Solarbio, China) following the manufacturer\u0026rsquo;s instructions. Gliding motility was examined as described by Bowman (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) and flexirubin-type pigment production was evaluated according to Bernardet and Bowman (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Growth under different pH conditions was assessed in marine broth (MB) adjusted to pH 4.0\u0026ndash;10.0 (in increments of 0.5 units) and pH 10.0\u0026ndash;12.0 (in increments of 1.0 unit) using the following buffers (50 mM each): MES (pH 4.0\u0026ndash;6.0), MOPS (pH 6.5\u0026ndash;7.0), Tris (pH 7.5\u0026ndash;8.5), CHES (pH 9.0\u0026ndash;10.0), and CAPS (pH 11.0\u0026ndash;12.0). NaCl tolerance was assessed in NaCl-free basal MB supplemented with 0\u0026ndash;20% NaCl (0\u0026ndash;4% at 0.5% intervals; 4\u0026ndash;20% at 1% intervals). The temperature range for growth was examined in MB at 4, 15, 20, 25, 28, 32, 35, 37 and 40\u0026deg;C, respectively. Anaerobic growth was evaluated on modified TYS broth (0.5% tryptone, 0.1% yeast extract, 3.0% sea salt) supplemented with 0.15% (w/v) cysteine hydrochloride and 1\u0026permil; (w/v) resazurin for 1 month in an anaerobic jar. Catalase activity was determined by bubble production in 3% (v/v) H\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e2\u003c/sub\u003e, and oxidase activity was assessed using commercial test strips (Merck). Hydrolysis of milk (0.5%, w/v), casein (0.5%, w/v), gelatin (1%, w/v), starch (1%, w/v), cellulose (1%, w/v), algin (1%, w/v) and Tweens 20, 40, 60, and 80 (1%, v/v) were carried out as described by Smibert and Krieg (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1981\u003c/span\u003e). Antibiotic susceptibility was tested by the disc diffusion method (Hudzicki \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) on MA plates using the following antibiotics: ampicillin (10 \u0026micro;g), cephalexin (30 \u0026micro;g), erythromycin (15 \u0026micro;g), roxithromycin (15 \u0026micro;g), oxytetracycline (30 \u0026micro;g), streptomycin (10 \u0026micro;g), vancomycin (30 \u0026micro;g), polymyxin B (300 IU), sulfadiazine (25 \u0026micro;g), sulfamethoxazole (25 \u0026micro;g), sulfafurazole (25 \u0026micro;g), norfloxacin (10 \u0026micro;g), levofloxacin (5 \u0026micro;g), lincomycin (2 \u0026micro;g) and chloramphenicol (30 \u0026micro;g). Additional enzyme activities and biochemical properties were examined by API ZYM and API 20NE test strips (bioM\u0026eacute;rieux), following standard protocols with a two-week extended incubation. Bacterial suspensions were prepared in 3% NaCl for API ZYM and in AUX medium for API 20NE, respectively.\u003c/p\u003e\n\u003ch3\u003eChemotaxonomic analysis\u003c/h3\u003e\n\u003cp\u003eFor cellular fatty acid analysis, the cells of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e and the reference strains were cultured in marine broth (MB) at 28\u0026deg;C and harvested during the early exponential growth phase (12\u0026ndash;24 h). Fatty acid methyl esters were prepared from lyophilized cells by saponification, methylation, and extraction following the standard protocol of the Sherlock Microbial Identification System (MIDI). They were then analysed using the Microbial Identification System (MIS) software for identification and quantification. Polar lipids and isoprenoid quinones were extracted from cells collected at the late exponential phase under the same culture conditions. Polar lipids were extracted according to the method of Komagata and Suzuki (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1988\u003c/span\u003e) and separated by two-dimensional thin-layer chromatography. Specific lipids were detected by spraying with the following reagents: phosphomolybdic acid (for total lipids), ninhydrin (for aminolipids), molybdenum blue (for phospholipids) (Collins and Jones \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1980\u003c/span\u003e). Respiratory quinones were analysed from lyophilized cells using a chloroform/methanol (2:1, v/v) extraction system. The extract was examined using a Dionex Ultimate 3000 HPLC system coupled to a Bruker impact HD mass spectrometer for identification.\u003c/p\u003e\n\u003ch3\u003eGenomic sequencing and analysis\u003c/h3\u003e\n\u003cp\u003eThe genome of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e was sequenced on an Illumina HiSeq X-ten platform (OE Biotech, Shanghai, PR China). Raw reads were quality-controlled using Trimmomatic and assembled using ABySS version 2.0. The assembled draft genome was submitted to the GenBank database. The DNA G\u0026thinsp;+\u0026thinsp;C content was calculated from the genome sequence. Genome relatedness was assessed by the average nucleotide identity (ANI) using the EzBioCloud tool (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ezbiocloud.net/tools/ani\u003c/span\u003e\u003cspan address=\"https://www.ezbiocloud.net/tools/ani\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) (Yoon et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and digital DNA-DNA hybridization (dDDH), estimated through the GGDC web server (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://ggdc.dsmz.de/ggdc.php\u003c/span\u003e\u003cspan address=\"http://ggdc.dsmz.de/ggdc.php\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) (Meier-Kolthoff et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).Functional annotation was performed using the RAST server (Aziz et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). To assess the polysaccharide-degrading potential, carbohydrate-active enzymes (CAZymes) were identified using the HMMER tool in the dbCAN3 server (Zheng et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Polysaccharide-utilization loci (PULs) were further predicted by searching the dbCAN-PUL database. Variation in glycoside hydrolase (GH) composition across different genera was visualised using non-metric multidimensional scaling (NMDS) based on Bray-Curtis dissimilarities.\u003c/p\u003e"},{"header":"Results and discussion","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e\u003cp\u003eThe nearly complete 16S rRNA gene sequence of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e (1491 bp; GenBank accession no. PV134465) was obtained in this study. Comparative analysis revealed that strain ASW11-125ᵀ belongs to the genus \u003cem\u003eChristiangramia\u003c/em\u003e, showing the highest sequence similarity to \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e (98.8%), followed by \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e (98.8%), \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e (98.6%), and other validly published type strains of this genus (93.3%\u0026ndash;98.3%). In the ML, NJ and ME phylogenetic trees (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, S1, and S2), strain ASW11-125\u003csup\u003eT\u003c/sup\u003e consistently formed a robust, distinct clade together with \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e, \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e and \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e. This subclade was stably positioned within a larger clade comprising all recognised species of the genus \u003cem\u003eChristiangramia\u003c/em\u003e, with strong bootstrap support (\u0026gt;\u0026thinsp;89%). Phylogenomic trees based on 1542 single-copy OCs confirmed that strain ASW11-125ᵀ, \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e and \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e formed a subclade, which further clustered with \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e into a larger clade (Figs.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, S3, and S4). Genomic relatedness analysis indicated that the average nucleotide identity (ANI) and digital DNA\u0026ndash;DNA hybridization (dDDH) values between strain ASW11-125ᵀ and the closely related strains were 91.5%, 85.2%, and 74.6% (ANI), and 44.3%, 28.7%, and 18.6% (dDDH), respectively (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). All values fell well below the accepted species demarcation thresholds (ANI\u0026thinsp;\u0026lt;\u0026thinsp;95\u0026ndash;96%; dDDH\u0026thinsp;\u0026lt;\u0026thinsp;70%), supporting the recognition of strain ASW11-125ᵀ as a novel species (Meier-Kolthoff et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Palmer et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003ePhenotypic and biochemical characterization\u003c/h3\u003e\n\u003cp\u003eStrain ASW11-125ᵀ was determined to be Gram-stain-negative, strictly aerobic, and capable of gliding motility. Flexirubin-type pigments were not produced. Transmission electron microscopy showed that the cells were short rods, approximately 0.5\u0026ndash;0.8 \u0026micro;m wide and 1.0\u0026ndash;3.0 \u0026micro;m long, and lacked flagella (Fig. S5). After 48 h of incubation on MA at 28\u0026deg;C, colonies were round, smooth, yellow, and 0.5\u0026ndash;1.0 mm in diameter. The strain grew at 4\u0026ndash;35\u0026deg;C (optimum 28\u0026deg;C), pH 6.0\u0026ndash;8.0 (optimum pH 7.0) and in the presence of 0.5\u0026ndash;16.0% NaCl (optimum 2.5\u0026ndash;3.0%). In antibiotic susceptibility tests, the isolate was resistant to ampicillin, cephalexin, erythromycin, streptomycin, sulfadiazine, and sulfafurazole, but susceptible to roxithromycin, oxytetracycline, sulfamethoxazole, norfloxacin, levofloxacin, vancomycin, polymyxin B, lincomycin, and chloramphenicol. A detailed comparison of the physiological and biochemical characteristics of strain ASW11-125ᵀ and its phylogenetic relatives is provided in the species description and in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eDifferential characteristics of strains ASW11-125\u003csup\u003eT\u003c/sup\u003e and type strains of closely related species\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCharacteristic\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGrowth range of:\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTemperature (optimum, ℃)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\u0026ndash;35 (28)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4\u0026ndash;37 (28\u0026ndash;30)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u0026ndash;36 (28\u0026ndash;30)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u0026ndash;40 (30)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4\u0026ndash;37 (23\u0026ndash;25)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e4\u0026ndash;40 (30)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNaCl (optimum, %)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.5\u0026ndash;16 (2.5\u0026ndash;3)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u0026ndash;15 (2\u0026ndash;5)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u0026ndash;6\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.5\u0026ndash;8 (1\u0026ndash;2)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1\u0026ndash;15 (4\u0026ndash;5)\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0\u0026ndash;9 (2)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003epH (optimum)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6.0\u0026ndash;8.0 (7.0)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.0\u0026ndash;10.0 (7.0\u0026ndash;8.0)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.0\u0026ndash;ND (7.0\u0026ndash;8.0)\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e5\u0026ndash;9 (7\u0026ndash;7.5)\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e6\u0026ndash;ND (7.0\u0026ndash;8.0)\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eAcid production from:\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGlucose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eUtilization of:\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eArginine\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eArabinose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMannose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMaltose\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eMalate\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eUrea\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eHydrolysis of:\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTween20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTween40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTween80\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eW\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStarch\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eEnzyme activity (API ZYM):\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLipase (C14)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTrypsin\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003ew\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eβ\u003c/em\u003e-Galactosidase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eα\u003c/em\u003e-Glucosidase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eN-acetyl glucosaminidase\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026ndash;\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e+\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e+\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eDNA G\u0026thinsp;+\u0026thinsp;C content (%)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e38.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e40.0\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e39.9\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e48.0\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e39.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e39.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"7\"\u003eStrains: 1, \u003cem\u003eC. qingdaonensis\u003c/em\u003e ASW11-125\u003csup\u003eT\u003c/sup\u003e; 2, \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e; 3, \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e; 4, \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e; 5, \u003cem\u003eC. echinicola\u003c/em\u003e KMM 6050\u003csup\u003eT\u003c/sup\u003e; 6, \u003cem\u003eC. sabulilitoris\u003c/em\u003e HSMS-1\u003csup\u003eT\u003c/sup\u003e.\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd colspan=\"7\"\u003eAll strains were Gram-negative and tested positive for catalase activities; hydrolysis of gelatin; and enzyme activities of alkaline phosphatase, esterase (C4), esterase lipase (C8), leucine arylamidase, valine arylamidase, cystine arylamidase, acid phosphatase and naphthol-AS-BI-phosphohydrolase. All strains were negative for citrate utilization. Data for columns 1\u0026ndash;4 were obtained in this study, while data for columns 5\u0026ndash;6 were taken from Nedashkovskaya et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2005\u003c/span\u003e), Park et al. (\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), respectively. +, positive; \u0026ndash;, negative; ND, no data available; W, weakly positive; *, Data from Li et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; \u0026dagger;, Data from Lau et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2005\u003c/span\u003e, Nedashkovskaya et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, Park et al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2015b\u003c/span\u003e.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003eChemotaxonomic characteristics\u003c/h2\u003e\u003cp\u003eThe major fatty acids (\u0026ge;\u0026thinsp;10%) of strain ASW11-125ᵀ were iso-C\u003csub\u003e15:0\u003c/sub\u003e (22.7%) and iso-C\u003csub\u003e17:0\u003c/sub\u003e 3-OH (10.0%), which are consistent with the general profile of the genus \u003cem\u003eChristiangramia\u003c/em\u003e. However, notable differences were observed in the proportions of several fatty acids compared to closely related strains. For example, strain ASW11-125ᵀ exhibited significantly lower proportion of C\u003csub\u003e16:0\u003c/sub\u003e (5.5%) than \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585ᵀ (11.5%). Similarly, its content of anteiso-C\u003csub\u003e15:0\u003c/sub\u003e (7.5%) was substantially lower than that in \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706ᵀ (14.8%) (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Polar lipid analysis indicated that strain ASW11-125\u003csup\u003eT\u003c/sup\u003e, along with \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e, \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e and \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e, all included phosphatidylethanolamine (PE) and two unidentified lipids (L1, L2). An additional unidentified lipid (L3) was detected in both ASW11-125ᵀ and \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585ᵀ, while an aminolipid (AL) was present only in \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366ᵀ (Fig. S6). The respiratory quinone of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e was MK-6, which aligns with the typical quinone system of the genus \u003cem\u003eChristiangramia\u003c/em\u003e.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eCellular fatty acid compositions (%) of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e and closely related strains of the genus \u003cem\u003eChristiangramia\u003c/em\u003e\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"5\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eFatty acid\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eStraight-chain\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e16:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e11.5\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e18:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eBranched\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e14:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e15:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e22.7\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e21.8\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e19.2\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e14.8\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e15:1\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e16:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e9.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e16:1\u003c/sub\u003e H\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eanteiso-C\u003csub\u003e15:0\u003c/sub\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e7.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e7.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u003cb\u003e14.8\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eHydroxy\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e15:0\u003c/sub\u003e 2-OH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e17:0\u003c/sub\u003e 2-OH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e3.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e3.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e15:0\u003c/sub\u003e 3-OH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e16:0\u003c/sub\u003e 3-OH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e4.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e2.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eiso-C\u003csub\u003e17:0\u003c/sub\u003e 3-OH\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e10.0\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e11.7\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e10.6\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eUnsaturated\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e15:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e6\u003cem\u003ec\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e0.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e17:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e6\u003cem\u003ec\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC\u003csub\u003e17:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e8\u003cem\u003ec\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e0.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eanteiso-C\u003csub\u003e17:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e9\u003cem\u003ec\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e2.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e1.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8.3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSummed features*\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e9.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9.3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e5.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e\u003cp\u003e6.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e5.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8.7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"5\"\u003eStrains: 1, \u003cem\u003eC. qingdaonensis\u003c/em\u003e ASW11-125\u003csup\u003eT\u003c/sup\u003e; 2, \u003cem\u003eC. marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e; 3, \u003cem\u003eC. portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e; 4, \u003cem\u003eC. aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e.\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd colspan=\"5\"\u003eAll data were obtained from this study. Major fatty acids in each strain are shown in bold.\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd colspan=\"5\"\u003e*Summed features are fatty acids that cannot be resolved reliably from another fatty acid using the chromatographic conditions chosen. The MIDI system reports these as a combined feature with a single percentage of the total; Summed feature 3 contains C\u003csub\u003e16:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e7\u003cem\u003ec\u003c/em\u003e and/or C\u003csub\u003e16:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e6\u003cem\u003ec\u003c/em\u003e and/or iso-C\u003csub\u003e15:0\u003c/sub\u003e 2-OH; Summed feature 9 contains iso-C\u003csub\u003e17:1\u003c/sub\u003e\u003cem\u003eω\u003c/em\u003e9\u003cem\u003ec\u003c/em\u003e and/or C\u003csub\u003e16:0\u003c/sub\u003e 10-methyl. ND, no data available.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eGenome features\u003c/h2\u003e\u003cp\u003eThe draft genome of strain ASW11-125\u003csup\u003eT\u003c/sup\u003e consists of 3,203,245 bp with 19 contigs, an N50 value of 880,254 bp, and a DNA G\u0026thinsp;+\u0026thinsp;C content of 38.3%. A total of 3,010 genes were predicted, including 2981 protein-coding genes, 6 pseudogenes, and 23 RNA genes (3 rRNA genes, 16 tRNA genes, and 4 other RNA genes) (Table S2). Genomic comparisons indicated that strain ASW11-125ᵀ had significantly higher gene abundances for amino acid/derivative metabolism (21.4%) and cofactor/vitamin/pigment metabolism (15.2%), but maintained comparable levels for protein (11.5%), carbohydrate (9.1%), and nucleoside/nucleotide (7.2%) metabolism relative to its close relatives (Table S3). Based on dbCAN3 analysis, 131 carbohydrate-active enzymes (CAZymes) were identified, accounting for 4.35% of the protein-coding sequences (Table S4). These included 68 glycosyltransferases (GTs, 16 families), 40 glycoside hydrolases (GHs, 25 families), 14 carbohydrate esterases (CEs, 4 families), 2 auxiliary activities (AAs, 2 families), and 7 carbohydrate-binding modules (CBMs, 6 families). NMDS analysis based on GH profiles revealed that the genus \u003cem\u003eChristiangramia\u003c/em\u003e clustered within a group of known polysaccharide-degrading \u003cem\u003eBacteroidota\u003c/em\u003e, including \u003cem\u003ePolaribacter\u003c/em\u003e, \u003cem\u003eFlavobacterium\u003c/em\u003e, \u003cem\u003eBacteroides\u003c/em\u003e, and \u003cem\u003eAlgibacter\u003c/em\u003e, and was clearly separated from the non-saccharolytic genus \u003cem\u003eStaphylococcus\u003c/em\u003e (ANOSIM, r\u0026thinsp;=\u0026thinsp;0.66, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.001; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea). Furthermore, \u003cem\u003eChristiangramia\u003c/em\u003e exhibited significantly greater similarity to \u003cem\u003eAlgibacter\u003c/em\u003e and \u003cem\u003ePolaribacter\u003c/em\u003e than to other genera (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.05) (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eb), indicating a closely aligned potential for polysaccharide degradation.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003ePolysaccharide-utilization loci for starch, laminarin and fructan\u003c/h2\u003e\u003cp\u003eStrain ASW11-125ᵀ harbors three polysaccharide utilization loci (PULs) that target starch, laminarin, and fructan. Each locus encodes a canonical PUL architecture, consisting of CAZymes, SusC/D-like transporters, and transcriptional regulators\u0026mdash;an arrangement that is conserved across \u003cem\u003eChristiangramia\u003c/em\u003e species (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Synteny across 18 \u003cem\u003eChristiangramia\u003c/em\u003e genomes showed the laminarin locus in every strain and starch loci in 94%, whereas the fructan locus is confined to one-third (33%) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). This distribution suggests that laminarin and starch are genus-wide metabolic pillars; fructan utilization constitutes a facultative trait likely maintained in specific ecological contexts. Collectively, these PULs deliver a rapid high-molecular-weight carbon uptake module that confers ecological plasticity and competitive supremacy throughout marine ecosystems.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003eTaxonomic conclusion\u003c/h2\u003e\u003cp\u003eThe phenotypic, phylogenetic, biochemical, genomic and chemotaxonomic features all support that strain ASW11-125\u003csup\u003eT\u003c/sup\u003e represents a novel species of the genus \u003cem\u003eChristiangramia\u003c/em\u003e, for which the name \u003cem\u003eChristiangramia qingdaonensis\u003c/em\u003e sp. nov. is proposed here.\u003c/p\u003e\u003cp\u003e\u003cb\u003eDescription of\u003c/b\u003e \u003cb\u003eChristiangramia qingdaonensis\u003c/b\u003e \u003cb\u003esp. nov.\u003c/b\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eChristiangramia qingdaonensis\u003c/b\u003e [qing.dao.nen\u0026rsquo;sis. N.L. fem. adj. qingdaonensis, pertaining to Qingdao, China, where the type strain was isolated]\u003c/p\u003e\u003cp\u003eCells are strictly aerobic, Gram-stain-negative, non-spore-forming, rod-shaped, non-flagellated and motile by gliding, measuring 0.5\u0026ndash;0.8 \u0026micro;m in width and 1.0\u0026ndash;3.0 \u0026micro;m in length. After 48 h on marine agar at 28\u0026deg;C, colonies are yellow, smooth, and 0.5\u0026ndash;1.0 mm in diameter. Growth occurs at 4\u0026ndash;35\u0026deg;C (optimally at 28\u0026deg;C), pH 6.0\u0026ndash;8.0 (optimally at pH 7.0) and in the presence of 0.5\u0026ndash;16.0% (w/v) NaCl (optimum 2.5\u0026ndash;3.0%). Flexirubin-type pigments are not produced. Cells are catalase-positive but oxidase-negative. Hydrolytic activities are observed for Tweens 20, 40, 60, and 80, starch, aesculin, casein, and gelatin, but not for cellulose or milk. Nitro-\u003cem\u003eβ\u003c/em\u003e-D-methylgalactose and malate are assimilated, while arabinose, mannose, mannitol, maltose, and citrate are not. Enzymatic activities are positive for alkaline phosphatase, esterase (C4), esterase lipase (C8), lipase (C14), leucine arylamidase, valine arylamidase, cysteine arylamidase, trypsin, chymotrypsin, acid phosphatase, naphthol-AS-BI-phosphohydrolase, \u003cem\u003eα\u003c/em\u003e-galactosidase, and \u003cem\u003eβ\u003c/em\u003e-glucosidase. However, activities are negative for urease, arginine dihydrolase, \u003cem\u003eβ\u003c/em\u003e-galactosidase, \u003cem\u003eα\u003c/em\u003e-glucosidase, and \u003cem\u003eβ\u003c/em\u003e-fucosidase. Acid production from glucose, nitrate reduction, and indole production are negative. The major polar lipid is phosphatidylethanolamine, and the predominant fatty acids are iso-C\u003csub\u003e15:0\u003c/sub\u003e (22.7%) and iso-C\u003csub\u003e17:0\u003c/sub\u003e 3-OH (10.0%). The respiratory quinone is menaquinone-6 (MK-6). The genomic DNA G\u0026thinsp;+\u0026thinsp;C content of the type strain is 38.3%. The type strain, ASW11-125\u003csup\u003eT\u003c/sup\u003e (=\u0026thinsp;KCTC 102340\u003csup\u003eT\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;MCCC 1K09555\u003csup\u003eT\u003c/sup\u003e), was isolated from intertidal sediment of Qingdao, PR China. The GenBank accession numbers for the 16S rRNA gene and whole-genome sequence are PV134465 and NZ_JBLWIM000000000, respectively.\u003c/p\u003e\u003c/div\u003e"},{"header":"Abbreviations","content":"\u003cdiv class=\"DefinitionList\"\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003eMCCC\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eMarine culture collection of China\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003eKCTC\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eKorean collection for type cultures\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003eMEGA\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eMolecular evolutionary genetics analysis\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003eRAST\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eRapid annotations using subsystems technology\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003eANI\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eAverage nucleotide identity\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv class=\"DefinitionListEntry\"\u003e\u003cdiv class=\"Term\"\u003edDDH\u003c/div\u003e\u003cdiv class=\"Description\"\u003e\u003cp\u003eDigital DNA-DNA hybridization\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003ch2\u003eCompeting interests\u003c/h2\u003e\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\u003c/p\u003e\u003cp\u003e\u003ch2\u003eEthics approval\u003c/h2\u003e\u003cp\u003eThis article does not contain any studies conducted by the authors involving human participants or animals.\u003c/p\u003e\u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e\u003cp\u003eThis work was funded by the National Natural Science Foundation of China (32300115).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eZTC designed the study, performed the data analysis, and wrote the manuscript. BY, YSL and ZWZ conducted genome analysis, contributed to the interpretation of data, and revised the manuscript. YL and XYZ supervised the experimental work, provided feedback on the manuscript, and contributed to the final manuscript revision.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eAll data obtained in this study are shown within the manuscript and supplemental materials.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAziz RK, Bartels D, Best AA, DeJongh M, Disz T, Edwards RA, Formsma K, Gerdes S, Glass EM, Kubal M, Meyer F, Olsen GJ, Olson R, Osterman AL, Overbeek RA, McNeil LK, Paarmann D, Paczian T, Parrello B, Pusch GD, Reich C, Stevens R, Vassieva O, Vonstein V, Wilke A, Zagnitko O (2008) The RAST Server: rapid annotations using subsystems technology. 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Microbiome 11:175. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1186/s40168-023-01618-7\u003c/span\u003e\u003cspan address=\"10.1186/s40168-023-01618-7\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"antonie-van-leeuwenhoek","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anto","sideBox":"Learn more about [Antonie van Leeuwenhoek](https://www.springer.com/journal/10482)","snPcode":"10482","submissionUrl":"https://submission.nature.com/new-submission/10482/3","title":"Antonie van Leeuwenhoek","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Christiangramia qingdaonensis, intertidal sediment, polyphasic taxonomy, polysaccharide utilization loci","lastPublishedDoi":"10.21203/rs.3.rs-8043186/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8043186/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eA novel polysaccharide-degrading bacterial strain, designated ASW11-125\u003csup\u003eT\u003c/sup\u003e, was isolated from intertidal sediments in Aoshan Bay, Qingdao, China. The strain was strictly aerobic, Gram-stain-negative, catalase-positive but oxidase-negative, short rod-shaped, and exhibited gliding motility without flagella. Growth occurred at 4\u0026ndash;35\u0026deg;C (optimum 28\u0026deg;C), pH 6.0\u0026ndash;8.0 (optimum pH 7.0), and in 0.5\u0026ndash;16.0% NaCl (optimum 2.5\u0026ndash;3.0%). The predominant polar lipid was phosphatidylethanolamine. Major fatty acids were iso-C\u003csub\u003e15:0\u003c/sub\u003e and iso-C\u003csub\u003e17:0\u003c/sub\u003e 3-OH, and the primary respiratory quinone was menaquinone-6 (MK-6). Based on the phylogenetic analyses of 16S rRNA gene sequences and 1542 single copy orthologous clusters, strain ASW11-125\u003csup\u003eT\u003c/sup\u003e affiliated with the genus \u003cem\u003eChristiangramia\u003c/em\u003e and was closely related to \u003cem\u003eChristiangramia portivictoriae\u003c/em\u003e MCCC 1A00585\u003csup\u003eT\u003c/sup\u003e (98.8%), \u003cem\u003eChristiangramia aquimixticola\u003c/em\u003e KCTC 42706\u003csup\u003eT\u003c/sup\u003e (98.8%) and \u003cem\u003eChristiangramia marina\u003c/em\u003e KCTC 12366\u003csup\u003eT\u003c/sup\u003e (98.6%). Comparative genomic analysis revealed that the average nucleotide identity (ANI) and digital DNA\u0026ndash;DNA hybridization (dDDH) values between strain ASW11-125ᵀ and its closely related species (74.6\u0026ndash;91.5% and 18.6\u0026ndash;44.3%, respectively) were clearly lower than the proposed species cutoff values. Based on a polyphasic characterization integrating phenotypic, phylogenetic, and chemotaxonomic evidence, strain ASW11-125ᵀ represents a novel species of the genus \u003cem\u003eChristiangramia\u003c/em\u003e, for which the name \u003cem\u003eChristiangramia qingdaonensis\u003c/em\u003e sp. nov. is proposed. The draft genome of strain ASW11-125ᵀ is 3.2 Mb in size with a G\u0026thinsp;+\u0026thinsp;C content of 38.3%. Notably, genomic analysis revealed an abundance of genes encoding putative carbohydrate-active enzymes (CAZymes), particularly those associated with starch, laminarin, and fructan utilization, suggesting its potential role in the marine carbon cycle. The type strain is ASW11-125ᵀ (=\u0026thinsp;KCTC 102340ᵀ = MCCC 1K09555ᵀ).\u003c/p\u003e","manuscriptTitle":"Christiangramia qingdaonensis sp. nov., a novel polysaccharide-degrading Bacteroidota bacterium, isolated from intertidal sediment","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-11-26 14:01:04","doi":"10.21203/rs.3.rs-8043186/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-12-02T02:15:38+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-12-01T07:04:05+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-11-22T02:27:15+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-11-21T23:46:51+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"168961591163543188300495851639414731549","date":"2025-11-21T00:14:10+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"105220304321221690672130544064605678357","date":"2025-11-20T05:46:17+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"10824455531188927723793792900815515464","date":"2025-11-18T00:10:26+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-11-17T02:16:36+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-11-07T14:24:33+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-11-07T04:46:00+00:00","index":"","fulltext":""},{"type":"submitted","content":"Antonie van Leeuwenhoek","date":"2025-11-06T03:16:38+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"antonie-van-leeuwenhoek","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"anto","sideBox":"Learn more about [Antonie van Leeuwenhoek](https://www.springer.com/journal/10482)","snPcode":"10482","submissionUrl":"https://submission.nature.com/new-submission/10482/3","title":"Antonie van Leeuwenhoek","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"830ff997-9ff9-4d57-b3ea-8aca6fb4bd17","owner":[],"postedDate":"November 26th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-01-12T16:12:47+00:00","versionOfRecord":{"articleIdentity":"rs-8043186","link":"https://doi.org/10.1007/s10482-025-02244-z","journal":{"identity":"antonie-van-leeuwenhoek","isVorOnly":false,"title":"Antonie van Leeuwenhoek"},"publishedOn":"2026-01-06 15:58:07","publishedOnDateReadable":"January 6th, 2026"},"versionCreatedAt":"2025-11-26 14:01:04","video":"","vorDoi":"10.1007/s10482-025-02244-z","vorDoiUrl":"https://doi.org/10.1007/s10482-025-02244-z","workflowStages":[]},"version":"v1","identity":"rs-8043186","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8043186","identity":"rs-8043186","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}