Comparative genomics of the Campylobacter genus: Insights into phylogenomics, virulence, genome plasticity and resistome profiling

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Several species, including C. jejuni, C. coli, C. fetus, C. concisus, C. lari, C. hyointestinalis, C. upsaliensis , and C. hepaticus , are established pathogens, while the pathogenic potential of other members remains unclear. This study presents a comparative genomics analysis of the fifty reported species of Campylobacter genus, encompassing phylogenomic relationships, functional repertoire profiling, virulence genes, diversity of Cytolethal distending toxin gene ( Cdt ), outer membrane components, genome plasticity, and resistome characterization. Phylogenetic analyses revealed that C. hepaticus, C. taniopygae , and C. iguaniorum , traditionally considered non-pathogenic or minor pathogens, cluster with major pathogenic species, suggesting shared evolutionary features. Functional repertoire profiling indicated metabolic flexibility that supports environmental adaptability, while virulence profiling highlighted both conserved and species-specific determinants. Variation in Cdt genes and outer membrane components emerged as key factors in pathogenicity. Notably, C. helveticus shows potential to emerge as a significant pathogen, whereas C. vicugnae and C. vulpis display close evolutionary relationships with C. jejuni . Genome plasticity analyses identified horizontal gene transfer via genomic islands, prophage insertions, and CRISPR arrays, underscoring the dynamic evolution of virulence traits. Resistome characterization revealed widespread antimicrobial resistance genes, raising concerns about multidrug resistance and clinical management. Overall, this study provides an integrative framework to understand the evolutionary dynamics, virulence potential, and antimicrobial resistance of Campylobacter , offering valuable insights for surveillance and therapeutic strategies. Campylobacter gastrointestinal infection cytolethal distending toxin cancer virulence horizontal gene transfer Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction Campylobacter is a genus of gram-negative bacteria belonging to the phylum Proteobacteria (Marroki et al. 2019). Campylobacter is slender, curved, spiral, motile, and non-spore-forming microbe (Igwaran et al. 2019). Multiflagella are present in some species such as C. showae , while in C. gracillis , flagella are absent (Acke 2018 ). Its shape is variable, ranging from 0.5 to 0.9 µm in length (Galanis 2007 ). The gastrointestinal tracts of different groups of animals are colonized by species of the Campylobacter genus (Gendy et al. 2013 ). Many species of the genus are associated with animals and humans and most commonly cause a diarrheal infection (Igwaran et al. 2019; Li 2018) known as campylobacteriosis. The most common cause of campylobacteriosis is consumption of contaminated poultry, although humans can also contract Campylobacter infections from ruminants including cattle, sheep, and goats (Kaakoush et al. 2015 ; Chlebicz and Śliżewska, 2018 ). Campylobacter is one of the most prevalent causative agents of food-borne infections in both developed and developing nations (Liu et al. 2022 ). Publications from 2014 to 2022 and the official public health websites of 195 nations declared that, Czech Republic had the highest prevalence of campylobacteriosis globally (215 per 100,000 in 2019, followed by Australia (146.8 per 100,000 in 2016) and New Zealand ( 126.1 per 100,000 in 2019) (Liu et al. 2022 ). According to a recent study conducted in Vellore, South India, utilizing PCR analysis on 400 human dysenteric stool samples collected between 2019 and 2020, Campylobacter was found to be the second most prevalent bacterial enteric pathogen (12%) (Lakshmi et al. 2022 ). Additionally, through the use of genus-specific PCR, Campylobacter spp., were detected in 16.77% of the samples collected from Periurban Bhubaneswar (Mohakhud et el. 2019). Another study performed on children under five in Northeast India between 2014 and 2015 found the prevalence of Campylobacter infection (10.1%), specifically C. jejuni (8.1%) in higher abundance (Borkakoty et al. 2020 ). In a very small number of cases (1 in 1000) C. jejuni also causes Gullian-Barre syndrome, an autoimmune neurological disease triggered by the molecular mimicry between outer membrane lipooligosaccharides and human peripheral nerve gangliosides (Li et al. 2020 ). Additionally, C. upsaliensis species of Campylobacter genus has been isolated from patients with bacteremia, hemolytic-uremic syndrome, spontaneous abortion, and Guillain-Barré syndrome (Wang et al. 2024 ). The genus Campylobacter has been divided into major and minor groups of pathogenic species (Rollins et al. 2006), with the major pathogenic species including C. jejuni, C. coli , and C. fetus and minor pathogenic species including C. concisus, C. lari, C. upsaliensis , C. hyointestinalis and C. hepaticus (Ienes et al. 2023). Chronic infections with Campylobacter spp. have also been observed to sometimes lead to colorectal cancer, Barrett's esophagus, and mucosa associated lymphoid tissue lymphoma (Marroki et al. 2019; Kato et al. 2021). Apart from the major and minor pathogenic species, some other members of the genus such as C. ureolyticus (Bullman et al. 2011 ; Bullman et al. 2012 ), C. helveticus, C. insulanigrae, C. mucosalis (Inglis et al. 2011 ), C. rectus (Lastovica et al. 2024) and C. sputorum have been associated with gastroenteritis (Lindblom et al. 1995 ; Man et al. 2011). Moreover, C. fetus has been shown to cause extra-intestinal infections and septic abortion in farm animals (Allos et al. 1995; Wu et al. 2022 ) Additionally, C. showae associated with crohns disease and ulcerative colitis (Man et al. 2011). A comparative genomic analysis of thirty-nine Campylobacter species has been conducted to date. This analysis identified antibiotic resistance genes, virulence factor genes, and pangenome genes. It also examined genetic diversity, evolutionary traits, and evaluated genome plasticity. Among the pathogenic species of Campylobacter, C. jejuni contributes for almost 90% of the instances of campylobacteriosis, while C. coli is responsible for less than ten percent of cases (Heimesaat et al. 2021 ; Liu et al. 2022 ). Numerous studies have linked prolonged gastrointestinal infections caused by C. jejuni to the development of colorectal cancer (Blaser et al. 2008; Guerra et al. 20011; He et al. 2019 ; Kato et al. 2023 ; Duijster et al. 2021 ) and mucosa associated lymphoid tissue (MALT) lymphoma (Louwen et al. 2013 ). C. jejuni produces a genotoxic substance known as cytolethal distending toxin (CDT), having DNAse activity (Lai et al. 2016 ; He et al. 2019 ; and Chen et al. 2024). CDT activity causes double-stranded DNA breaks and increases the probability of transformation of normal cells into cancer cells (Brauner et al. 2010 ; Vogtmann et al. 2016; Tremblay et al. 2021 ). Apart from CDT, various other virulence factors are involved in the establishment and continuation of infection by C. jejuni (Kemper et al. 2023). These virulence factors are related with bacterial adherence to intestinal mucosa, flagella mediated motility, invasive capability, chemotaxis, quorum sensing and outer membrane component lipopolysacchaides (LPS). Even though a number of species of this genus have been identified as pathogenic (major or minor), the pathogenic potential of most of the species has not yet been investigated. This study was designed with the aim of elucidating the phylogenomic relationships between different species of the genus Campylobacter and the analysis of their pathogenic potential by the comparative analysis of their virulence genes, antibiotic resistance genes and metabolic pathways. Apart from this, we have identified the genomic islands; prophage regions and CRISPRs analysis in order to determine the genomic plasticity among the species of Campylobacter genus. Methods Phylogenomic analysis To perform phylogenomic analysis, the genomes of fifty Campylobacter species whose genome sequences were publicly available were downloaded from the NCBI database (Table 1 ). Phylogenetic analysis based on conventional 16s rRNA gene sequences was performed, for which the RNAmmer version 1.2 server was used to extract 16s rRNA genes from each genome (Lagesen et al. 2007 ). A phylogenetic tree was constructed using the maximum likelihood algorithm (Felsenstein et al. 1993) with a 1000 bootstrap value in MEGA X. To assure the accuracy of the phylogenetic relationship multiple conserved bacterial marker protein based phylogenetic analysis was performed, for which 31 conserved bacterial marker proteins were extracted from each of the 50 species of Campylobacter using AmphoraNet server (Kerepesi et al. 2014 ). A phylogenetic tree was constructed using the Neighbor Joining algorithm (Saitou et al.1987) with 1000 bootstrap revaluations implemented in MEGA X (Erickson et al. 2014 ). Further, whole genome data-based phylogenomic analysis was performed, which includes Average nucleotide identity (ANI) and tetranucleotide frequency. ANI values were calculated by employing the BLASTALL algorithm of J species server version 1.2.1. (Richter et al. 2009). To calculate ANI values and tetra nucleotide frequency, genomes from each species were uploaded to J species server version 1.2.1. After that two-way pair matrix was formed containing pairwise ANI scores. Hierarchial clustering was performed and dendogram was constructed using MeV 2.0 software for ANI and tetranucleotides frequency scores. Table 1 Accession numbers, isolation sources and general genomic features of Campylobacter species. Species name Strain NCBI accession number Source of isolation Genome size (in Mb GC Content (%) Number of contings Plasmid tRNAs Number of coding sequences Campylobacter jejuni Gundogdu et al. 2007 ATCC 70089 NC_002163.1 Not known 1.6 30.5 1 Absent 44 1687 C. armoricus Miller WG; Yee 2020 (direct submission) CCUG 73571 NZ_CP053825.1 River water France: Brittany 1.6 28.5 1 Absent 46 1625 C. avium Miller 2017(direct submission) LMG 24591 NZ_CP022347.1 Caecal host-chicken Italy: Bolognan 1.7 34.2 1 Absent 40 1783 C. concisus Miller WG 2016 (direct submission) ATCC 33237 NZ_CP012541.1 Gingival sulcus host-homo sapiens USA 1.8 37.5 1 Absent 47 1898 C. coragiensis Miller WG 2020 (direct submission) LMG27932 NZ_CP053842.1 Feces host-Macaca silenus Ireland: Cork 1.7 31.9 2 38 Kb (NZ_CP053843.1) 41 1767 C. coli Kane et al. 2017(direct submission) Aerotolerant OR12 NZ_CP019977.1 Organic chicken farm United Kingdom: Lincolnshire 2 30.8 1 Absent 44 2196 C. curvus Miller WG 2020 (direct submission) ATCC 35224 NZ_CP053826.1 Human, jaw abscess USA: Massachusetts 2 44.5 1 Absent 48 2010 C. hominis Fouts 2007 (direct submission) ATCC BAA-381 NC_009714.1 Not known 1.7 31.5 2 36 Kb (NC_009713.1) 44 1738 C. fetus Davis et al. 2021 CF00A031 NZ_CP059443.1 Preputial wash Host –bovine Canada:British Columbia 1.8 33.5 2 26 Kb (NZ_CP059444.1) 44 1811 C. geochelonis (direct submission 2016) RC7 NZ_FIZQ01000001.1 Cloacal swab 2 33.5 34 ND 41 2070 C. gracilis Miller et al. 2015 ATCC 33236 NZ_CP012196.1 Oral Host-Homo sapiens USA 2.3 46.5 1 Absent 43 2757 C. helveticus Miller WG 2017 (direct submission) ATCC 51209 NZ_CP020478.1 Feces Host-cat Switzerland-Berne 1.9 34.5 6 1. 72 Kb (NZ_CP020479.1) 2. 34 Kb(NZ_CP020480.1) 3. 32 Kb (NZ_CP020481.1) 4. 16 Kb(NZ_CP020482.1) 5. 17 Kb ( NZ_CP020483.1) 42 2028 C. hepaticus Van et al. 2019 (direct submission) HV10 NZ_CP031611.1 Gallus gallus (whole organism) Australia, Victoria 1.5 28 1 Absent 43 1519 C. hyointestinalis Miller WG 2020 (Direct submission) Lawsoni CHY5 NZ_CP053828.1 Procrine stomatch 1.8 33.5 2 44 Kb (NZ_CP053829.1) 44 1880 C. iguaniorum Gilbert et al. 2016 (Direct submission) 2463D NZ_CP010995.1 Iguana iguana Netherlands 1.8 35.5 1 Absent 44 1895 C. insulaenigrae Miller et al. 2014 (direct submission) NTCC 12927 NZ_CP007770.1 Marine mammal UK Scotland 1.5 28 1 Absent 42 1485 C. lanienae Miller et al. 2017 NCTC 13004 NZ_CP015578.1 Feces host-homo sapiens Switzerland 1.6 34.5 1 Absent 40 1634 C. lari Miller et al. 2014 RM 2100 NC_012039.1 Not known 1.6 29.5 3 1. 46 Kb (NC_012040.1) 2. 18 Kb ( CP000933.1) 46 C. Mucosalis Miller WG 2020 (direct submission) ATCC 43264 NZ_CP053831.1 Pig, small intestine UK 1.8 36.5 1 Absent 43 1834 C. ornithocola Miller WG 2020 (direct submission) LMG 29815 NZ_CP053848.1 Wild bird fecal samples Chile: Valdivia 1.6 29.4 1 Absent 46 1627 C.pinnipediorum Miller WG 2016 (direct submission) RM172661 NZ_CP012547.1 Sea lion lung USA: California 1.7 30.5 1 Absent 45 1775 C. rectus Miller WG 2016 (direct submission) ATCC 33238 NZ_CP012543.1 Oral host-homosapiens, USA: Boston 2.6 44.5 1 Absent 48 3083 C. showae Miller WG 2016 (direct submission) ATCC 51146 NZ_CP012544.1 Oral host-homosapiens, Japan: Showa 2.1 45.5 1 Absent 47 2312 C. novaezeelandiae Bloomfield; Wilkinson D.A. 2020 (direct submission) B423b NZ_QPGR01000001.1 Anas platyrhynchos, New Zealand 1.6 27.5 59 ND 39 1645 C. sputorum Miller et al. 2017 LMG 11764 NZ_CP019684.1 Human, fecal Canada: Ottawa 1.7 29.5 1 Absent 47 1765 C. upsaliensis Miller 2020 (direct submission) RM3940 NZ_CP053849.1 Human, fecal USA: Los Angeles, California 1.6 35 1 Absent 44 1668 C. ureolyticus Kyrpides et al. 2013 (direct submission) DSM 20703 NZ_KB894730.1 Amniotic fluid, Canada 1.7 29 34 ND 41 1803 C. volucris Miller et al. 2014 LMG 24379 NZ_CP007774.1 Black headed gull, Sweden 1.5 28.5 1 Absent 43 1520 C. subantarcticus Miller et al.2014 LMG 24377 NZ_CP007773.1 Grey headed albatross, South Georgia and the South Sandwich Islands 1.9 29.8 1 Absent 46 1918 C. vulpis Parisi 2015. (direct submission) 73/13 NZ_LDWY01000065.1 Fox blind gut, Italy: Monterenzio 1.6 34.5 95 ND 34 1667 C. protucalensis Silva et al. 2020 (direct submission) FMV-PI01 VWSJ01000001.1 Bovine preputial sample, Portugal: Lisbon 1.7 28.3 98 ND 39 1798 Candidates C. infans Duim 2021 ( direct submission) 19S00001 NZ_CP049075.1 Feces Host-homosapiens, Netherlands: Utrecht 1.8 35.9 2 5.8 Kb (NZ_CP049076.1) 38 1871 C. taeniopygiae Mannion 2018 (direct submission) MIT10-5678 NZ_NXLY01000001.1 Feces Host- Taeniopygia guttata, USA 1.7 29.5 88 ND 40 1824 C. estrildidarum Mannion 2018 (direct submission) MIT17-664 NZ_NXLZ01000001.1 Feces Host- Taeniopygia guttata, USA 1.7 29.2 48 ND 42 1798 C. aviculae Mannion 2018 (direct submission) MIT17-670 NZ_NXMA01000001.1 Feces Host- Taeniopygia guttata, USA 1.7 29.5 59 ND 42 1752 C. canadensis Miller WG 2019 (direct submission) LMG 24001 NZ_CP035946.1 cloacal swab host- Whooping crane Canada: Calgary 1.9 27.5 1 Absent 41 1885 C. troglodytis Mannion A 2018 (direct submission) MIT 05-9149A NZ_QHLI01000001.1 Chimpanzees, USA: MIT Cambridge, MA 2.9 35 295 ND 41 4083 C. blaseri Miller WG 2020 (direct submission) LMG 30333 NZ_CP053841.1 Feces Host- Phoca vitulina, Netherlands:Pieterburen 1.9 29.5 1 Absent 42 1887 C. pleoridis Lane 2020 (direct submission) 2016D-0074 NZ_CP063079.1 Shellfish 1.7 28.5 2 45 Kb (NZ_CP063080.1) 46 1665 C. cuniculorum Miller et al. 2017 (direct submission). DSM 23162 = LMG 24588 NZ_CP020867.1 Rabbit, caecal, Italy: Bologna 1.9 31.2 3 1. 49 Kb (NZ_CP020868.1) 2. 18 Kb (NZ_CP020869.1) 40 1996 C. analoticus Aydin et al. 2021 faydin-G140 NZ_JAGSSY010000001.1 Faeces of Anatolian Ground squirrel in Turkey 1.8 35.2% 13 ND 43 1968 C. bilis Phung et al. 2022 VicNov18 WUED01000001.1 Chikens with spotty liver disease 1.4 30.7 41 ND 41 1568 C. cryaerophila Neill et al. 1985 LMG 24291 NXGK01000077.1 Bovine Brain, aborted fetus, Ireland 2 27.2 91 ND 40 2145 C. devanensis Miller et al. 2017 (direct submission) NCTC 13003 NZ_CP018788.1 Unkown 1.6 33.5 1 Absent 40 1618 C. magnus Gruntar et al. 2023 46386 JAQSLK010000001.1 Caecal contents of domestic pigs. 1.7 38.8 19 ND 38 1862 C. majalis Lynch et al. 2022 LMG 7974 CAJHOF010000001.1 Procelli gastro-intestinal mucosa 2.2 33.8 59 ND 39 2086 C. massiliensis Antezacker et al. 2021 Marseille-Q3452 JACLZK010000001.1 Dental source host-homo sapiens, France 2.2 45.7 3 ND 47 2452 C. procelli Miller et al. 2024 RM6137 CP018789.1 Wild pig, fecal USA: California 1.6 34.2 2 ND 40 1674 C. suis Lynch et al. 2022 LMG 8286 CAJHOE010000001.1 Procrine gastro-intestinal mucosa 1.7 37.2 16 ND 40 1827 C. vicugnae Miller et al. 2024 RM12175 CP018793.1 Alapacs goats and sheep 1.6 32.4 3 ND 40 1669 ND*-not determined Functional potential analysis Functional flexibility among Campylobacter spp. was determined by comparative genomic analysis, which is based on metabolic pathways. For this, amino acid sequences of all the fifty species of Campylobacte r were retrieved from the RAST server (Aziz et al. 2008 ), followed by the gene finding using KAAS (KEGG Automatic Annotation Server) by BLAST against the KEGG (Kyoto Encyclopedia of Genes and Genomes) database (Ogata et al. 1999 ). Metabolic pathways were reconstructed using MinPath (Minimal Set of Pathways) (Ye et al. 2009). One-way hierarchical clustering was performed based on 125 metabolic pathways. For hierarchical clustering, a heat map was created using the Euclidian and Ward algorithms, followed by a matrix transformation of all values with Log 10 and keeping 500 of the rows with the highest standard deviation (Ryan et al. 2019 ). Analysis of the genes associated with virulence factors Previous studies have identified the presence of virulence factors in Campylobacter genus (Igwaran et al. 2019; Kemper et al. 2023; Bunduruș et al. 2023 ). These virulence factors were identified in the genomes of different species of this genus, using the annotation generated by RAST and a heat map was generated by the NGCHM builder (Ryan et al. 2019 ). Cytolethal distending toxin (Cdt) is a key virulence factor found in Campylobacter species with potential link to colorectal cancer (Kato et al. 2023 ). As a result, Cdt genes were examined in each species within the genus. Species containing Cdt genes were subjected to a BLASTn analysis (Altschul et al. 1997 ) against the Cdt genes of C. jejuni to assess the similarity of Cdt genes from other Campylobacter species to Cdt gene of C. jejuni. For this analysis subunit A, B and C of each species concatenated and compared. Further, a multiple sequence alignment of Cdt protein of subunit B sequences was conducted using Muscle (see Fig. 5 ) to identify the conserved domains within the protein. A phylogenetic tree was constructed based on the amino acid sequence (concatenated sequence of subunit A, B and C) of the Cdt gene using the Neighbor-Joining method in Mega X software. Type IV ((T4SS) and Type VI secretion systems(T6SS) play a crucial role in host-pathogen interactions and are present in several well-known bacterial pathogens, including Salmonella, Pseudomonas, Yersinia , and Vibrio (Pukatzki et al. 2007 ). Consequently, proteins associated with Type IV and Type VI secretion systems have been identified in each Campylobacter species. The outer membrane components of Campylobacter serve as multifunctional virulence factors, playing a crucial role in the bacterium's ability to colonize, invade, and survive within the human host (Lopes et al. 2021 ). To classify lipopolysaccharides (LPS) from Campylobacter species based on their components, we examined the structures of lipid A, the outer core, and the inner core in each species of the genus. These were categorized into rough LPS, deep rough LPS, and smooth-type LPS. Additionally, sialyated carbohydrate lipooligosaccharide (LOS) structures synthesis enzymes on the outer membranes of the members of this genus were identified as they have been known to be associated with complement-mediated nerve fatalities and severe colitis in C. jejuni (Louwen et al. 2013 ). Further, LOS components identified were compared to the LOS of C. jejuni to assess their similarity. Genome plasticity and resistome analysis Genomic islands, along with other mobile and accessory genetic elements, play a significant role in the horizontal movement of some genes in bacterial populations and contribute to the genomic flexibility of bacteria (Dobrindt et al. 2004 ). Genomic islands were identified using Genomic Island Viewer 4 (Bertelli et al. 2017 ) based on variations in GC content and codon usage bias and IspandPath-DIMOB. The presence of prophage regions within bacterial genomes can signify the occurrence of horizontal gene transfer in bacteria (Borodovich et al. 2022 ). The detection of prophage regions in Campylobacter species was conducted using the PHASTER server (Arndt et al. 2016 ). Subsequently, to ascertain which genes in Campylobacter species were acquired through phages, bacterial gene sequences integrated with prophage regions were identified. Furthermore, CRISPR genes were extracted from each genome by uploading the genomes to the CRISPRFinder online server (Grissa et al. 2007 ). This server conducted a BLAST search against the dbCRISPR database, categorizing the results into true and false CRISPRs based on their association with CRISPR-associated (cas) genes. Additionally, the spacer sequences of Campylobacter species were subjected to a BLAST search against the NCBI virus database to identify similarities with viral sequences, thereby predicting which viruses may be involved in horizontal gene transfer within Campylobacter species. Furthermore, antibiotic resistance genes were identified in the fifty species of Campylobacter genus using the resistance gene identifier 6.0.1 (Kwatra 2021 ). Results Phylogenomic analysis A general analysis of the genomic characteristics of 50 Campylobacter species indicated that their genome sizes ranged from 1.4 Mb to 2.9 Mb, with GC content between 27.2% and 46.5%. The species and genomes examined in this study were predominantly isolated from food sources, including meat, poultry, and animal-associated environments. Among the 50 species of the genus, plasmids were identified in nine species, with plasmid sizes ranging from 16 Kb to 58 Kb (Table 1 ). Plasmid encoded genes associated with type IV secretion system in C. fetus, C. pleoridis, C. helveticus and C. lari species. Phylogenetic analysis based on the 16S rRNA genes revealed that C. jejuni and C. coli , which are major pathogenic species, form a monophyletic clade. In addition, C. fetus (major pathogenic species) and C. iguaniorum (non-pathogenic species) formed a monophyletic clade. However, C. cryaerophila is distinct among 50 spp. of the Campylobacter . To further elucidate the phylogenetic relationships among species within the genus, phylogenetic analyses were conducted using 31 conserved bacterial marker genes. These analyses revealed that C. coli, C. hepaticus , and C. jejuni spp. are the most closely related species, forming a monophyletic clade. Additionally, other species, including C. helveticus, C. upsaliensis (minor pathogenic species), and C. vulpis , are closely related and grouped within a single clade, while C. fetus, C. iguaniorum , and C. hyointestinalis constitute another monophyletic clade. ANI provides strong resolution for genomes that share 80–100% ANI, indicating they belong to the same species, as well as for closely related species that share less than 80% ANI. In the present study, the ANI values ranged from 68 to 72% among Campylobacter spp. Whole-genome analyses utilizing ANI and TETRA frequency metrics have demonstrated that C. jejuni, C. coli, C. hepaticus , and C. taniopygae are species with close phylogenetic relationships (Fig. 1 ). Comparative analysis of functional repertoire of Campylobacter genus Reconstruction of metabolic pathways for the 50 species within the Campylobacter genus, based on KAAS, led to the identification of several conserved metabolic pathways, including glycolysis, the tricarboxylic acid (TCA) cycle, the pentose phosphate pathway, pentose and glucuronate interconversion, fructose and mannose metabolism, and the galactose metabolism pathway. Notably, the fatty acid degradation pathway was absent in Candidatus C. infans, C. laninae , and C. avium . Additionally, certain pathways were absent in specific species; for instance, the histidine metabolism pathway was present in all species except C. taeniopygae . The benzoate degradation pathway was absent in C. curvus but present in all other species. Furthermore, the beta-alanine metabolic pathway was not found in C. taeniopygae, C. aviculae , and C. canadensis . With the exception of C. hominis , all Campylobacter species exhibited pathways for dioxin and xylene degradation, biofilm formation ( Pseudomonas aeruginosa ), and unsaturated fatty acid biosynthesis. Moreover, the glucose inotate biosynthesis pathway was absent in C. novazeelandiae and C. taeniopygae . Similarly, the mineral absorption pathway was exhibited by all species but was absent in C. blaseri . Ascorbate and validomycin biosynthesis pathways were present in C. coli, C. geochelonsis, C. helveticus, C. insulanigare , and C. troglodytis , but absent in other species. Some metabolic pathways associated with the pathogenicity of the bacterium and antibiotic resistance were found to be present in all species of the genus, including the bacterial secretion system, bacterial chemotaxis, quorum sensing, two-component system, lipopolysaccharide synthesis, and biofilm formation E. coli , platinum drug resistance, antifolate resistance, cationic antimicrobial peptide resistance vanomycin resistance and beta-lactum resistance. Hierarchical clustering of the metabolic pathways across the 50 genomes revealed a close relationship among C. jejuni, C. coli, C. hepaticus , and C. bilis , as corroborated by phylogenomic analysis (Fig. 2). Figure 2 The heat map illustrates the comparative functional abundance of metabolic pathways among Campylobacter species. The clustering of species with C. jejuni , based on their metabolic pathways, is emphasized, including C. coli, C. hepaticus , and C. bilis . The threshold of the number of genes associated with specific pathways is represented through a color scale Comparative analysis of factors associated with virulence (I) Genes associated with virulence Virulence factors impart specific traits associated with virulence, including bacterial adherence to the intestinal mucosa, motility mediated by flagella, and the ability to invade host tissues. Genes involved in imparting virulence (n = 37) that were identified in Campylobacter genus included chemotaxis virulence factors CheA, CheB, CheR, CheV, CheW, CheZ Cet A, Cet B, LuxS, motility virulence factors JlpA, flaB, flaA, Flagellin C, major outer membrane protein, outer membrane protein CadF, adhesion protein A CapA, periplasmic binding protein PEB1A, fibronectin-like protein A FlpA, type IV secretion system, type VI secretion system, fibronectin F, racR, dnaJ, docA, flagellin C, Campyloabcter invasion antigen B CiaB, invasion antigen C, Campylobacter invasion antigen CiaD, invasion associated protein iamA, periplasmic protein HtrA, expression of invasion pldA, sialyltransferase, intracellular invasion Cial, Lipopolysaacharide production WlaN, Sialyltransferase, methyl accepting chemotaxis protein, and cytolethal distending toxin. Species clustering on the basis of these factors demonstrated the close associatoion of C. jejuni , with C. vicugnae and C. coli with C. rectus (Fig. 3 ). Among the major pathogenic species of the Campylobacter genus, twenty nine, twenty six and twenty four virulence factors have been identified in C. jejuni, C. coli and C. fetus respectively. Additionally, among the minor pathogenic species, nineteen, twenty two, twenty five, twenty six and twenty virulence factors were identified in C. concisus , C. hyointestinalis , C. lari , C. upsaliensis and C. hepaticus respectively. High number of virulence factor genes were also identified in species of the genus, which have not been classified as major or minor pathogens but have been reported to be associated with gastroenteritis, including C. heleveticus (n = 29), C. insulanigare (n = 24) and C. mucosalis (n = 26), C. sputorum (n = 22), C. rectus (n = 20), C. ureolyticus (n = 10) and C. showae (= 22) have been associated with colitis. Furthermore, high number of virulence factor genes (n ≥ 19) were also identified in species which have not been reported as pathogenic till date including C. armoricus, C. avium, C. iguaniorum, C. laninae, C. ornithocola, C. pinnipediorum, C. novaezeelandiae, C. subantracticus, C. volucris, C. taeniopygae, C. estrildidarum, C. troglodytis, C. vulpis, C. aviculae, C. pleridids , C. cuniculorum , and C. procelli and C. vicugnae. Previous studies have indicated the association of cytolethal distending toxin ( Cdt ) of Campylobacter jejuni with initiation of colon cancer in patients suffering from chronic infections (He et al. 2019 ; He et al. 2024 ; Guidi. 2014; Graillot et al. 2016 ; Hartl et al. 2020; Yusuf et al. 2023 ). In this study, we identified and compared the Cdt genes across 50 Campylobacter species. This analysis revealed that 27 out of the 50 species harbor Cdt genes, specifically C. jejuni, C. coli , and C. fetus (major pathogens); C. lari, C. upsaliensis and C. hyointestinalis (minor pathogens); C. helveticus, C. insulanigrae and C. mucosalis (associated with gastroenteritis); C. vulpis, C. taeniopygae, C. armoricus, C. aviculae, C. estrildidarum, C. subantracticus, C. pleoridis, C. volucris, C. geochelonsis, C. iguaniorum, C. laninae,, C. devanensis, C. procelli, C avium, C. ornithocola, C. canadensis, C. vicugnae and C. bilis (pathogenicity has not been reported yet). Cdt consists of three subunits: A, B and C, with subunit B being associated with DNase activity (Brauner et al. 2010 ). The gene encoding for subunit B of the Cdt gene of C. jejuni is composed of 798 bp in size, the gene for subunit A consists of 807 bp, and that of subunit C is made up of 570 bp. Genes encoding for all three subunits were identified in 22 species. The gene for subunit C was not identified in C. iguaniorum , C. canadensis , and that for subunit A was not located in C. avium, C. vicugnae and C. devanensis indicating non-functional Cdt proteins in these species. The BLASTn analysis of Cdt nucleotide sequences of 22 Campylobacter species (in which all three subunits were present) against the Cdt gene of C. jejuni revealed that the Cdt gene of C. bilis exhibited highest similarity with Cdt gene of C. jejuni with 74.05% identity e-value 2e-75) however, its nucleotide sequences of B subunit was truncated (size of subunit B of C. bilis was 573 bp). Furthermore, C. vulpis exhibited the highest similarity to the Cdt gene of C. jejuni (71.70% identity; e-value 0.0) followed by C. helveticus (71.34% identity; e-value 0.0) and C. upsaliensis (70.78% identity; e-value 0.0). The identity percentage of the Cdt genes of other species with that of C. jejuni ranged from 69.53% to 64.57% (Fig. 4 ). Furthermore, multiple sequence alignment of the amino acid sequences of Cdt genes from Campylobacter species revealed the conserved regions on catalytic sites of subunit B that may be critical for the function of the Cdt protein across different species including, H152, D185 and D222 ( D222 conserved in all spp. except C. bilis ) (Fig. 5 ). The phylogenetic tree based on protein sequences of the Cdt gene demonstrated the C. jejuni formed a distinct cluster, separated from the other Campylobacter species. The distinct clustering pattern indicates that Cdt gene in C. jejuni has undergone unique evolutionary pressure, which may contribute to its well- established pathogenic potential compared to other members of the genus (Fig. 6 ). Both T4SS and T6SS play important roles in pathogenicity (Costa et al. 2019). Analysis of T4SS proteins in each species of the Campylobacter genus revealed that T4SS proteins were present in 29 out of the 50 species. However, not all species included the complete set of system proteins (Fig. 7 ). Examination of the T6SS genes in Campylobacter species led to their identification in 21 out of the 50 species (supplementary table 1 online resource). Generally; 13–15 core genes form the cluster encoding T6SS. (II) Outer membrane pathogenic component analysis The lipopolysaccharides that were identified among the species of the Campylobacter genus were classified based on their components into rough LPS (lipid A and outer core), deep rough LPS (lipid A and inner core), and smooth LPS (complete LPS: lipid A, outer core, inner core, and O-antigen). Among the species of the genus, 41 species have deep rough LPS, making them highly susceptible to hydrophobic antibiotics. C. jejuni, C. armoricus, C. laninae, C. hyointestinalis, C. upsaliensis, C. bilis, C. cryaerophila, C. devanensis and C. vicugnae were found to have rough LPS, making it less susceptible to antibiotics (Table 2 ). Furthermore, sialylated oligosaccharide structures synthesis enzymes including ADP-heptose-lipooligosaccharide heptosyl transferase, CMP-Neu5Ac synthase and CMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase were examined in each species of the Campylobacter genus to assess their potential for pathogenicity based on their membrane components. Forty one species were found to harbour the sialylated lipooligosaccharide synthesis enzyme known as ADP-heptose-lipooligosaccharide heptosyl transferase. This finding suggests that presence of sialyated oligosaccharide structure in these forty one species may have incorporated in their virulence. Additionally, the BLAST result of ADP-heptose-lipooligosaccharide heptosyl transferase of forty species against C. jejuni ADP-heptose-lipooligosaccharide heptosyl transferase exhibited that, out of forty species, only four species showed a significant identity percentage, including C. coli , which showed 83% (e-value 0.0) identity; C. taniopygae , which showed 78.5% ( e-value 2e-162) identity; C. ornithocola , which represented 77.47% (e-value 4e-74) identity; and C. lari , which showed 76.71% (e-value 4e-69) identity. However, thirty six species exhibited no significant similarity with C. jejuni ADP-heptose-lipooligosaccharide heptosyl transferase. Apart from this, CMP-Neu5Ac synthase was found to be present in thirty species and CMP-N-acetylneuraminate-beta-galactosamide-alpha-2, 3-sialyltransferase was present in thirty two species. Table 2 Types of lipopolysaccharides and lipooligosaccharide synthesis enzymes in Campylobacter species S.No. Campylobacter species Lipid A Outer core Inner core O-antigen Type of lipopolysaccharides (LPS) ADP-heptose-lipooligosaccharide heptosyl transferase CMP-Neu5Ac synthase CMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase 1 C. jejuni Present Present Present Absent Rough LPS Present Present Present 2 C. coli Present Absent Present Absent Deep rough LPS Present Present Present 3 C. fetus Present Absent Present Absent Deep rough LPS Present Present Absent 4 C. armoricus Present present Present Absent Rough LPS Present Present Present 5 C. aviculae Present Absent Present Absent Deep rough LPS Present Absent Present 6 C. blaseri Present Absent Present Absent Deep rough LPS Present Present Absent 7 C. canadensis Pressent Absent Present Absent Deep rough LPS Present Present Present 8 C. cuniculorum Present Absent Present Absent Deep rough LPS Present Absent Present 9 C. curvus Present Absent Present Absent Deep rough LPS Present Absent Present 10 C. estrildidarum Present Absent Present Absent Deep rough LPS Present Absent Present 11 C. gracillis Present Absent Present Absent Deep rough LPS Absent Absent Absent 12 C. helveticus Present Absent Present Absent Deep rough LPS Present Present Present 13 C. hyointestinalis Present Present Present Absent Rough LPS Absent Absent Present 14 C. mucosalis Present Absent Present Absent Deep rough LPS Present Present Present 15 C. Pinnipediorum Present Absent Present Absent Deep rough LPS Present Present Absent 16 C. portucalensis Present Absent Present Absent Deep rough LPS Present Present Present 17 C. showae Present Absent Present Absent Deep rough LPS Present Absent Absent 18 C. sputorum Present Absent Present Absent Deep rough LPS Absent Absent Absent 19 C. subantracticus Present Absent Present Absent Deep rough LPS Present Absent Present 20 C. troglodytis Present Absent Present Absent Deep rough LPS Present Absent Absent 21 C. upsaliensis Present Present Present Absent Rough LPS Present Present Present 22 C. ureolyticus Present Absent Present Absent Deep rough LPS Absent Present Present 23 C. volucris Present Absent Present Absent Deep rough LPS Present Absent Present 24 C. vulpis Present Absent Present Absent Deep rough LPS Present Present Present 25 C. taeniopygae Present Absent Present Absent Deep rough LPS Present Present Present 26 C. avium Present Absent Present Absent Deep rough LPS Present Present Present 27 C. concisus Present Absent Present Absent Deep rough LPS Present Absent Absent 28 C. corcagiensis Present Absent Present Absent Deep rough LPS Present Absent Absent 29 C. geochelonsis Present Absent Present Absent Deep rough LPS Absent Absent Absent 30 C. hepaticus Present Absent Present Absent Deep rough LPS Present Present Present 31 C. hominis Present Absent Present Absent Deep rough LPS Absent Absent Absent 32 C. iguaniorum Present Absent Present Absent Deep rough LPS Absent Present Absent 33 C. insulanigrae Present Absent Present Absent Deep rough LPS Present Present Present 34 C. laninae Present Present Present Absent Rough LPS Absent Present Present 35 C. lari Present Absent Present Absent Deep rough LPS Present Absent Present 36 C. novaezeelandiae Present Absent Present Absent Deep rough LPS Present Present Present 37 C. ornithocola Present Absent Present Absent Deep rough LPS Present Present Present 38 C. pleoridis Present Absent Present Absent Deep rough LPS Present Absent Present 39 C. rectus Present Absent Present Absent Deep rough LPS Present Absent Absent 40 Candidates C. infans Present Absent Present Absent Deep rough LPS Present Present Absent 41 C. analoticus Present Absent Present Absent Deep rough LPS Absent Present Absent 42 C. bilis Present Present Present Absent Rough LPS Present Present Present 43 C. cryaerophila Present Present Present Absent Rough LPS Present Present Absent 44 C. devanensis Present Present Present Absent Rough LPS Present Present Present 45 C. magnus Present Absent Present Absent Deep rough LPS Present Present Absent 46 C. majalis Present Absent Present Absent Deep rough LPS Absent Present Present 47 C. massiliensis Present Absent Present Absent Deep rough LPS Present Present Absent 48 C. procelli Present Absent Present Absent Deep rough LPS Present Present Present 49 C. suis Present Absent Present Absent Deep rough LPS Present Absent Absent 50 C. vicugnae Present Present Present Absent Rough LPS Present Present Present Genome plasticity analysis Horizontal gene transfer (HGT) is a critical mechanism in the evolution of microbial genomes. In numerous bacterial pathogens, genomic islands and other mobile genetic elements facilitate movement of genetic repertoire and have been implicated in genome evolution via HGT (Juhas et al. 2015). Consequently, genomic islands were identified in twenty-nine species including, C. armoricus, C. coli, C. avium, C. blaseri, C. canadensis, C. consisus, C. corcagiensis, C. cuniculorum, C. curvus, C. fetus, C. gracillis, C. helveticus, C. hominis, C. hyointestinalis, C. iguaniorum, C. laninae, C. lari, C. mucosalis, C. ornithocola, C. pinnipediorum, C. rectus, C. showae, C. sputorum, C. subantracticus, C. upsaliensis, Candidatus C. infans, C. procelli and C. vicugnae . Certain genes associated with virulence were mapped to these genomic islands. These findings suggest that some virulence factors in these species may have been acquired through HGT (Table 3 ). Table 3 Genes associated with virulence and pathogenicity mapped to the genomic islands among Campylobacter spp. Species Genes mapped to the genomic islands C. fetus Type II toxin-antitoxin system, YafQ family toxin and type II toxin-antitoxin system RaE/ParE C. coli Type VI secretion system protein, Type VI secretion system associated protein, type II CRISPR RNA guided endonuclease Cas9 C. subantracticus Type VI secretion system associated lipoprotein, Type VI secretion lysozyme-like protein C. iguaniorum LPS biosynthesis protein, toxin-antitoxin system protein. C. gracilis host nuclease inhibitor protein Gam, pathogenicity locus C. rectus DNA type IV secretion system protein ComB1a, type IV secretary system conjugative DNA transfer family protein, CRISPR associated protein, type VI secretion system C. laninae TetM/ TetW/ Tet O/ Tet S family tetracycline resistance ribosomal protection protein. C. procelli Cytolethal distending toxin CdtB C. helveticus Type IV secretion system and cytolethal distending toxin subunit A, B C. cuniculorum Type IV secretion system protein virB9 was present. C. avium Type II secretion system protein. C. canadensis Type II secretion system protein. C. hyointestinalis Type IV secretary system conjugative C. blaseri Type IV secretion system DNA-binding domain-containing protein, virulence protein. Candidates C. infans Type II and type III secretion system protein, type I restriction endonuclease, type IV secretion system, virulence protein. It has been established that prophages play key role in HGT (Mancini et al. 2024 , Gomathinayagam et al. 2025). Among the 50 species of Campylobacter , prophage regions were identified in 27 species, out of which 9 species harbored intact prophages (complete detail for supplementary table 2 (online resource) and Table 4 ). Most of the genes incorporated in the prophage region were annotated as hypothetical. Table 4 Genomic islands, prophages, crisper arrays and spacers among the spp. of Campyloabacter Species Genomic Islands Intact prophages Crisper arrays Spacers C. jejuni - - 1 1 C. coli 30 2 4 9 C. fetus 89 - 2 44 C. armoricus 12 - 1 1 C. aviculae - - - - C. blaseri 6 1 1 8 C. canadensis 4 - 4 42 C. cuniculorum 6 - 1 29 C. curvus 4 - 2 15 C. estrildidarum - - - - C. gracillis 8 2 3 95 C. helveticus 6 - - - C. hyointestinalis 21 - - - C. mucosalis 2 - - - C. Pinnipediorum 8 1 - - C. portucalensis - - 1 1 C. showae 6 - - - C. sputorum 2 - - - C. subantracticus 36 1 - - C. troglodytis - - - - C. upsaliensis 2 - - - C. ureolyticus - - - - C. volucris - - - - C. vulpis - - - - C. taeniopygae - - - - C. avium 2 - - - C. concisus 6 1 - - C. corcagiensis 4 1 - - C. geochelonsis - - 1 78 C. hepaticus - - - - C. hominis 8 1 1 71 C. iguaniorum 20 1 1 7 C. insulanigrae - - - - C. laninae 4 - 1 18 C. lari 10 - - - C. novaezeelandiae - - - - C. ornithocola 6 - - - C. pleoridis - 1 11 C. rectus 36 1 4 100 Candidates C. infans 4 - 1 22 C. analoticus - - 1 1 C. bilis - - 1 3 C. cryaerophila - - 1 1 C. devanensis - - 5 9 C. magnus - - 9 10 C. majalis - - 1 1 C. massiliensis - - 2 46 C. procelli 12 - 5 6 C. suis - - - - C. vicugnae 4 - 2 20 The CRISPR-Cas system is an adaptive immune system in bacteria and archaea that captures and stores sequences from invading elements (phages, plasmids) as spacers. Out of 50 species, twenty-six species were found to harbor CRISPR arrays (complete detail supplementary table 3 (online resource)) and Table 4 . A total of seven hundred spacers were identified in Campylobacter spp., which denotes frequent infection by bacteriophages. Identification of spacers on the viral database revealed the prevalence of Podoviridae and Myoviridae as the major phages infecting Campylobacter species. Resistome analysis Antibiotic resistance genes in Campylobacter species pose a substantial public health challenge, as these bacteria are frequently implicated in foodborne illnesses. The presence of such genes complicates therapeutic interventions and may lead to increased morbidity among affected individuals. An analysis of antibiotic resistance genes across 50 Campylobacter species revealed that 28 species possess genes conferring resistance to antibiotics (Table 5 ). However, antibiotic resistance genes were not identified in 22 species including C. fetus and C. hepaticus which are pathogenic. Table 5 Antibiotic resistance genes present in Campylobacter species. Species Antibiotic resistance drug class AMR gene Family Resistance mechanism C. jejuni, C. coli, C. lari, C. volucris, Candidates C. infans, C. magnus Penicillin beta-lactam, macrolide antibiotic, fluoroquinolone antibiotic, cephalosporin, fusidane antibiotic OXA beta-lactamase, OXA-61-like beta-lactamase, resistance-nodulation-cell division (RND) antibiotic efflux pump Antibiotic inactivation C. portucalensis, C. armoricus, C. avium, C. blaseri, C. concisus, C. corcagiensis, C. geochelonsis, C. hyointrstinalis, C. ornithocola, C. pleoridis, C. rectus, C. showae, C. subantracticus, C. ureolyticus, C. analoticus, C. cryaerophila, C. devanensis, C. majalis, C. massiliensis, C. vicugnae. Fluoroquinolone antibiotic, tetracycline antibiotic, glycopeptide antibiotic Resistance-nodulation-cell division (RND) antibiotic efflux pump, glycopeptide resistance gene cluster, vanT Antibiotic efflux, antibiotic target alteration C. magnus, C. taeniopygae, C. laninae Tetracycline antibiotic Tetracycline-resistant ribosomal protection protein Antibiotic target protection C. taeniopygae Aminoglycoside antibiotic APH(3') Antibiotic inactivation Discussion Comparative genomic analysis of prokaryotes has enhanced our understanding of the biology of various pathogenic microorganisms. The Campylobacter genus includes a group of bacteria known for their significant role in causing infections in both humans and animals (Igwaran & Okoh, 2019 ). Several Campylobacter species have been known to cause serious infections in humans, underscoring their clinical significance. Over the last decade, the global incidence of campylobacteriosis has risen, indicating that it has become a widespread infectious disease (Bukari et al. 2025 ). In Campylobacter species, variations in genome size and the presence of plasmids are crucial factors that influence their adaptability, pathogenicity, and resistance to antimicrobials. Most Campylobacter spp. have a relatively small genome, typically ranging between 1.4 to 2.9 Mb, indicating their streamlined metabolic capabilities and reliance on host environments. Despite its compact genome, Campylobacter displays significant genetic diversity owing to frequent horizontal gene transfers and recombination. Plasmids were found to be harbored by nine spp. of Campylobacter in this study, which greatly contribute to this diversity by carrying genes linked to virulence. For example, genes on plasmids of C. fetus, C. pleoridis, C. helveticus and C. lari encoding type IV secretion systems can enhance survival in challenging environments and promote gene exchange between species. Apart from that, the gene encoding cag pathogenicity island protein (cag12) was annotated on the plasmids of C. pleoridis, C. helveticus and C. lari . Therefore, genome size and plasmid content are one of the factors associated with the determination of the ecological fitness of Campylobacter species. The phylogenomic relationships elucidated in this study offer a comprehensive understanding of the connections between pathogenic and non-pathogenic species within the genus, revealing that pathogenic species are related to some nonpathogenic or minor pathogenic species. Phylogenetic and phylogenomic analyses of Campylobacter species have shown intriguing clustering patterns, where major pathogenic species, such as C. jejuni and C. coli , are closely grouped with minor or non-pathogenic species. This close association underscores the high degree of genetic relatedness within the genus and suggests that differences in pathogenicity of the spp. may stem from subtle variations in gene content, mobile genetic elements, and regulatory mechanisms, rather than broad genomic divergence. The clustering of pathogenic species with minor pathogenic members, such as C. hepaticus , C. upsaliensis or C. hyointestinalis , indicates that the evolutionary paths of Campylobacter are shaped by gene acquisition and loss events, horizontal gene transfer, and niche-specific adaptations. Comparative genome analysis based on metabolic pathways demonstrated a close relationship among C. coli, C. bilis , and C. jejuni , indicating that these species share a conserved metabolic framework that supports their survival and adaptation in diverse host environments. Clustering with C. bilis , in which pathogenicity has not been reported, yet, highlights possibility that core metabolic pathways are conserved across pathogenic and non-pathogenic members of the genus. The close metabolic relatedness highlights a common evolutionary origin and functional similarity in nutrient utilization and energy metabolism, however, the absence of pathogenic traits in C. bilis despite metabolic similarity to C. jejuni and C. coli underscores the importance of virulence repertoires in pathogenicity of the species rather than metabolic capacity alone. Even though the species of Campylobacter have been established as major or minor pathogens and been implicated in gastrointestinal infections, however, the information regarding the pathogenic potential of the rest of the species remains scarce. In this study, a genome-wide exploration of the factors associated with virulence was performed which provided key insights into virulence potential of non-pathogenic species. Based on the virulence repertoire, C. vicugnae was clustered with C. jejuni. C. vicugnae has been recently isolated from a patient suffering from gastroenteritis (Jehanne et al. 2024 ), however, its pathogenicity has not been established yet. This indicates that C. vicugnae harbors a subset of virulence associated determinants despite its limited association with clinical infections which may suggest either the presence of cryptic or unexplored virulence potential in C. vicugnae , or an evolutionary retention of virulence genes that are not actively expressed under natural conditions. Overall, this observation highlights the need for further functional and epidemiological studies to assess whether C. vicugnae could act as an opportunistic pathogen under certain conditions, or whether its virulence gene repertoire primarily represents evolutionary remnants without significant pathogenic outcomes. Some virulence factors associated with chemotaxis, adhesion and invasion play very important roles in the bacterial pathogenicity. High number of virulence factors (n ≥ 20) were observed in non pathogenic species (apart from major and minor pathogenic species) including C. armoricus, C. avium, C. iguaniorum, C. laninae, C. ornithocola, C. showae, C. novaezeelandiae, C. volucris, C. subantracticus, C. taeniopygae, C. estrildidarum, C. aviculae, C. troglodytis, C. devanensis, C. vicugnae, C. vulpis, C. pleoridis, C. cuniculorum, C. procelli and C. helveticus, C. insulanigare, C. mucosalis, C. rectus, C. sputorum , which have been associated with gastroenteritis and which have never been reported as pathogenic. The highest number of genes for virulence factors (n = 29) were harbored by C. jejuni , and C. helveticus . C. helveticus has been linked to gastroenteritis; however it has not been classified as a major or minor pathogen. The enrichment of adhesion, invasion, and toxin related genes in C . jejuni is consistent with its well established role as the leading cause of human campylobacteriosis, while the presence of similar virulence gene repertoire in C. helveticus highlights its emerging significance as a potential pathogen. The higher virulence gene load may provide the species carrying them with a broader arsenal to overcome host defense mechanisms, establish persistent colonization, and cause disease. Such findings also indicate possible selective pressure or horizontal gene transfer events that have shaped the virulence architecture of these species. The process of chemotaxis involves the detection of external signals by bacteria, facilitating their movement towards favorable environments. This mechanism is employed by several pathogenic bacteria to invade their hosts. In Campylobacter species, specifically in C. jejuni , chemotaxis is mediated by glycoproteins and mucin, which serve as chemoattractants, guiding the bacteria to primary localization sites within the avian gut and the mucus-filled crypts of the avian ceca (Chang and Miller 2006 ). The chemotaxis process utilizes two-component signal transduction pathways, involving six proteins: chemotaxis A, B, R, W, Y, and Z (Hermans et al. 2012), along with two methyl-accepting proteins. The CheW protein functions as a coupling protein, linking methyl-accepting proteins to CheA, which subsequently activates and transfers the phosphoryl group to either CheY or CheB. The binding of CheY proteins to the flagellar motor component FliM induces a change in rotation from counterclockwise to clockwise, thereby influencing bacterial swimming and motility. In addition to the six chemotaxis proteins, other proteins such as CetA, CetB, and LuxS are also implicated in the chemotaxis process in Campylobacter species. Regarding adhesion as a virulence factor, Krause Craszczynska et al (2007) have previously reported that the CadF gene is present in all C. jejuni and C. coli strains, is an outer membrane protein facilitating cell adhesion through its binding to the cell adhesion protein fibronectin. In present study, the CadF gene was identified in 44 out of 50 Campylobacter species. The colonization of Campylobacter species in the host gut necessitates adherence to host gastrointestinal epithelial cells (Jin et al. 2001 ). Glycoproteins binding to fibronectin activate signaling GTPases Rac1 and Cdc42, which promote Campylobacter cell internalization (Ziprin et al. 1999 ). In terms of invasion, bacterial flagella play a crucial role during host invasion, serving as export apparatuses for the secretion of non-flagellar proteins (Guerry et al. 2007). The Flac and Cia gene products utilize the flagellar secretion apparatus to be delivered into the host cell cytoplasm, which is essential for colonization and invasion into the host (Carrillo et al. 2004 ). Among the 50 species of Campylobacter , invasion genes CiaB and HtrA were identified 48 and 44 species respectively indicating invasion potential in these species. The cancer promoting potential of C. jejuni in prolonged infections has been established and has been attributed to the cytolethal distending toxin produced by the species (Marroki et al. 2019; Zergui et al. 2025 ). The Cdt complex is composed of three subunits: A, B, and C. Thereby, all three subunits of Cdt genes were annotated in 22 species in this study. Apart from this, incomplete or non-functional Cdt genes missing either subunit A or subunit C genes were identified in C. iguaniorum and C. canadensis (no subunit C) and C. avium, C. devanensis and C. vicugnae (no subunit A). As per previous study, Cdt A and Cdt C attach to cholesterol-rich micro-domains on the cell membrane, facilitating the entry of Cdt B into cells via clathrin-coated pit endocytosis (Nesic et al. 2005). Once internalized Cdt B moves to the cytosol and may be transported retrogradely from the Golgi complex to the endoplasmic reticulum. Eventually, the Cdt B relocates to the nucleus, guided by a potential nuclear localization signal in its amino acid sequence. In the nucleus, Cdt B causes DNA double-strand breaks, leading to cell-cycle arrest (Lai et al. 2016 ). Thus, Cdt can be considered as an important virulence marker as well as a key factor for promotion of tumor formation. The gene encoding Cdt was identified in all major (n = 3) and 3 minor pathogenic species (out of 5) of the genus as well as 3 species (out of 6) which have been associated with gastroenteritis. However, the Cdt gene was also identified in 13 species in which pathogenicity has not been reported yet. Comparative analysis of the gene encoding Cdt in different Campylobacter species with Cdt encoding gene of C. jejuni , revealed that Cdt gene of C. vulpis (non-pathogenic) exhibit a high similarity to the Cdt gene of C. jejuni . Furthermore, multiple sequence alignment of the amino acid sequences of Cdt B protein among different Campylobacter species revealed conserved and variable regions that may contribute to their functional roles. The conservation of key amino acids such as histidine H152 and aspartate D185, D222 (except C bilis ) on catalytic sites across Cdt B protein sequences suggests evolutionary pressure to maintain structural and enzymatic function essential for Cdt activity. The presence of Cdt ABC gene in species not typically associated with disease suggests that possession of some virulence genes alone does not necessarily translate into virulence potential and may reflect an evolutionary remnant retained from a common ancestor or may have been acquired through horizontal gene transfer. Acquisition of other virulence factors in future might provide them with a complete arsenal for virulence. Phylogenetic analysis of the concatenated amino acid sequences of Cdt ABC revealed that C. jejuni constitutes a distinct cluster, indicating unique sequence variations in comparison to other Campylobacter species. This distinct separation highlights the evolutionary divergence of Cdt in C. jejuni , which may contribute to its notable pathogenic potential. However, aside from the distinct clustering of C. jejuni ; C. helveticus, C. upsaliensis , and C. vulpis are closely related to C. jejuni based on the amino acid sequences of Cdt , indicates a conserved evolutionary pattern and might have potential functional similarity of the toxin among these species. The secretion systems are essential for bacterial pathogenesis, adaptation, and interactions between host and pathogen. In Campylobacter species, the T4SS and T6SS have become significant molecular mechanisms that enhance virulence, facilitate genetic exchange, and ensure survival in various environments. These systems, which are specialized protein complexes, enable bacteria to introduce effector proteins into eukaryotic (host cells) and prokaryotic (other bacteria) target cells. This injection can disrupt biological functions, enhance bacterial survival and reproduction within the host, and exacerbate disease (Wang et al. 2021 ). Many bacteria, particularly gram-negative bacteria, possess the complex machinery known as T4SS. A set of VirB/VirD genes or their homologs, initially identified in Agrobacterium tumefaciens , encode T4SS. T4SS typically includes an extracellular pilin composed of a minor VirB5 subunit and a major VirB2 subunit. Substrate secretion is driven by three ATPases, VirB4, VirB11, and VirD4, which may also assist in system assembly. The inner membrane channel comprises the biotopic membrane proteins VirB8 and VirB10, as well as the polytopic membrane protein VirB6, according to biochemical and functional studies. The composition of the pore at the outer membrane, which allows the substrate to enter the extracellular milieu, remains unknown. The short lipoprotein VirB7 may be a component of the complex VirB9. However, neither protein possesses a transmembrane region that has been identified or predicted. Additionally, the roles of VirB1 and VirB3 in this complex remain unclear (Fronzes et al. 2009 ). The T4SS genes were found on the plasmid in some species of the genus, including C. fetus, C. lari, C. helveticus , and C. pleoridis , among which which C. pleoridis is a non-pathogenic species. Their plasmid encodes many proteins of T4SS, indicating that the plasmid-associated secretion system may play a role beyond virulence, potentially contributing to ecological fitness rather than direct pathogenicity. Among the fifty species of Campylobacter , T4SS proteins were identified in 29, indicating a likelihood of their involvement in horizontal gene transfer, plasmid-related functions, and adaptation. However, the uneven presence of the T4SS proteins among species indicates differing functionality and evolutionary diversification within the genus. Moreover, as reported by Gabbert et al ( 2023 ), the T4SS was absent in C. jejuni . The impact of the absence of T4SS is limited to its reduced capacity to perform functions typically associated with classical T4SS, such as horizontal gene transfer via conjugation or the direct injection of effectors into host cells, as seen in other pathogens. Instead, C. jejuni has adapted to utilize other secretion systems, specifically the Type III secretion system (T3SS) and the T6SS, to facilitate its pathogenesis (Tikhomirova et al. 2024 ). T4SS proteins were identified in a subset of species, suggesting that this system is not universally conserved throughout the genus. Apart from that, generally, 13–15 core genes form the cluster encoding T6SS. This system mimics an inverted phage tail by puncturing target cells with a contraction-based mechanism. The Hcp gene, linked to the T6SS, creates a tube-like structure that allows certain substances and effector carriers to be secreted. The protein VgrG resembles a spike and penetrates the target membrane to deliver effectors. ClpV is an ATPase necessary for the recycling of sheath components such as tssC . The sheath components are tssB , which contracts during secretion, and tssC , which pairs with tssB . The base plate component tssE anchors the sheath. The baseplate core assembly genes are tssH, tssG , and tssK . The tssH gene is associated with the disassembly of the sheath for recycling (Pisarz et al. 2024 ). A membrane complex protein gene includes tssJ, tssL , and tssM . Our analysis of T6SS across 50 Campylobacter species reveals insights into the distribution patterns of these systems. Among the 50 species, 21 contained T6SS proteins, with 5 being major or minor pathogens, 2 linked to gastroenteritis, and 14 species for which pathogenicity has not yet been reported. Conversely, T6SS is increasingly recognized as a widespread and versatile secretion system, contributing to both inter-bacterial competition and environmental adaptation. Our study indicates that T6SS proteins are unevenly distributed among different Campylobacter species. Previous research has established that the lipopolysaccharides constituting the outer membrane components of gram-negative bacteria, are well-documented virulence factors due to their impermeability to various toxins and enzymes, including proteases, lysozyme, detergents, and some hydrophobic antibacterial agents (Matsuura et al. 2013; Zhao et al. 2016). LPS is composed of three primary parts: lipid A, the core region, and O-antigen (O-polysaccharide). LPS is released in large amounts as a result of the lysis of bacterial cells, which can become an endotoxin, leading to pathological conditions, including sepsis. Thus, LPS is a well-established bacterial virulence factor responsible for pathogenesis (Sidor et al. 2023). Prior studies have shown that deep-rough LPS is more susceptible to antibiotics and reduce pathogenicity by obstructing the ADP heptose pathway (Sidor et al. 2023). An analysis of the outer membrane lipopolysaccharides of 50 Campylobacter species revealed that 41 species exhibited and deep rough LPS and 9 species exhibited rough LPS which is more resistant to antibiotics. Furthermore, the sialylated Foligosaccharide structure found on C. jejuni species is known to enhance their potential for pathogenicity. These structures possess epitopes that closely resemble ganglioside epitopes present in human peripheral nerves. C. jejuni is classified into two categories based on whether the sialic acid component is linked to the bacterial carbohydrate outer surface through the action of sialyltransferases Cst-II or Cst-III. These bacteria exhibit a highly pathogenic phenotype and possess the capability to cause severe colitis (Louwen et al. 2013 ). The sialyated lipooligosaccharide synthesis enzymes analysis indicates that enzyme ADP-heptose lipooligosaccharide (LOS) heptosyltransferase, found in Campylobacter species, plays a vital role in the synthesis and structural alteration of LOS, a crucial element of the outer membrane. This enzyme facilitates the addition of heptose units from ADP-heptose to the developing LOS core, thereby enhancing the structural variety and functional intricacy of LOS across various Campylobacter species. These alterations are linked to immune system evasion, resistance to serum, and interactions between the host and pathogen. Notably, differences in heptosyltransferase genes between pathogenic and non-pathogenic Campylobacter species may determine the degree of LOS truncation or extension, thereby influencing bacterial virulence potential. The presence of the enzyme in forty one species of the genus indicates evolutionary preservation of LOS biosynthesis pathways, while species-specific variations may signify adaptive strategies to unique ecological niches or host environments. Apart from that, the detection of CMP-Neu5Ac synthase in Campylobacter species underscores a crucial element of their sialic acid metabolism. In present study CMP-Neu5Ac synthase was found to be present in thirty species. This enzyme facilitates the activation of N-acetylneuraminic acid (Neu5Ac) by transforming it into CMP-Neu5Ac, which acts as a donor substrate for sialyltransferases that modify surface structures like lipooligosaccharides (LOS). In C. jejuni and some related species, CMP-Neu5Ac synthase is pivotal in promoting the sialylation of LOS, a mechanism that imitates host gangliosides, aiding in immune evasion, persistence, and occasionally leading to post-infectious neuropathies such as Guillain–Barré syndrome (Pérez et al. 2018). Notably, the presence and distribution of CMP-Neu5Ac synthase differ among Campylobacter species, with pathogenic strains more commonly possessing this gene than non-pathogenic species. This variation implies that acquiring and maintaining CMP-Neu5Ac synthase may offer a selective advantage in host-associated environments by boosting virulence and adaptation. Consequently, the presence of CMP-Neu5Ac synthase not only highlights metabolic diversity within the genus but also signifies its evolutionary significance in shaping host-pathogen interactions. Furthermore, the enzyme CMP-N-acetylneuraminate β-galactosamide α-2,3-sialyltransferase, found in thirty two species of Campylobacter , is a vital component of the sialylation process, connecting sialic acid metabolism to alterations in surface structures. This enzyme uses CMP-Neu5Ac as a donor to attach sialic acid to the terminal galactose units of lipooligosaccharides (LOS), resulting in sialylated LOS formation. These changes are crucial for host-pathogen interactions, as they allow bacteria to mimic human gangliosides, aiding in immune system evasion and resistance to serum. In pathogenic strains such as C. jejuni , the function of this sialyltransferase is closely linked to increased virulence, survival within the host, and the potential to cause post-infectious autoimmune conditions such as Guillain–Barré and Miller Fisher syndromes. However, this enzyme is not consistently present across the genus; many non-pathogenic or less virulent species do not possess it. This uneven distribution indicates that the presence and maintenance of CMP-N-acetylneuraminate β-galactosamide α-2, 3-sialyltransferase offers a selective benefit specifically for pathogenic Campylobacter , highlighting its role in adaptive evolution and host colonization. Horizontal gene transfer (HGT) is a pivotal evolutionary mechanism in prokaryotic genomes, facilitating the exchange of genetic material between organisms that are not in a parent-offspring relationship. This process is a well-documented adaptation strategy observed in bacteria and archaea. Although the implications of HGT extend beyond pathogenic organisms, it is frequently associated with microbial antibiotic resistance and pathogenicity (Soucy et al. 2015 ). A substantial component of HGT is mediated by genomic islands (GEIs), which are distinct DNA segments that vary among closely related strains. Some GEIs are mobile, while others are either immobile or have lost their mobility. These islands can integrate into the host chromosome, excise, and transfer to a new host through transformation, conjugation, or transduction. GEIs are vital in the evolution of diverse bacterial species, as they facilitate the dissemination of variable genes, including those conferring antibiotic resistance and virulence (Juhas et al. 2009 ). Genomic islands were detected in 29 Campylobacter species. The genomic islands in most species of this genus contain genes crucial for pathogenicity, such as pathogenic loci, type IV and type VI secretion systems, toxin-antitoxin systems, genes associated with antibiotic resistance, virulence genes, and cytolethal distending toxin subunits B and A. Toxin-antitoxin system in Campylobacter is genetic elements that help to maintain plasmid stability and contributes to bacterial survival under stress. This suggests that HGT might be an important source for the evolution of pathogenic potential in this genus. Additionally, analysis of mobile genetic elements revealed that prophage regions were present in 27 species of Campylobacter , with 9 species harboring intact prophages. CRISPR arrays were detected in 25 species, comprising a total of 700 spacers, suggesting their potential role in regulating HGT and shaping genomic plasticity within the genus. Furthermore, analysis of antibiotic resistance genes revealed that most pathogenic species exhibit these genes, while some nonpathogenic species show only 50% or less similarity with antibiotic resistance genes, indicating that pathogenic species have developed adaptive potential against drugs. The ability of microorganisms to acquire antibiotic resistance is well-established. The overuse and indiscriminate application of clinically prescribed antibiotics, coupled with the ongoing development and spread of mobile genetic resistance elements, have led to the emergence of multidrug-resistant (MDR) and even extremely drug-resistant (XDR) bacterial pathogens over the past two decades. The most commonly used antimicrobials for treating campylobacteriosis, when clinical intervention is necessary, are fluoroquinolones and macrolides; however, these are becoming less effective against Campylobacter . Antibiotic resistance genes were found in 28 out of 50 species of the genus, indicating a broad but uneven distribution of resistance within the group. The species that possess these resistance genes demonstrate their ability to withstand antimicrobial pressure, raising concerns for treatment options. Conclusion Campylobacter represents a clinically and epidemiologically important genus. Some species of the genus are key enteric pathogens. Comparative genomics of the Campylobacter genus provides comprehensive insights into its evolutionary complexity, pathogenic potential and adaptive strategies. Phylogenetic and phylogenomic analysis showed that several non-pathogenic or minor pathogenic species of Campylobacter clustered with major pathogenic species, indicating shared genomic features that could confer potential for pathogenicity under certain evolutionary or environmental conditions. Functional repertoire profiling underscored the metabolic flexibility of the genus, whereas virulence gene analysis, including detailed analysis of the Cdt gene, indicated that C. helveticus possesses the potential to emerge as a pathogen. C. vicugnae clustered with C. jejuni suggesting shared virulence traits though its pathogenicity not yet well established, and the analysis of Cdt gene revealed, high similarity between C. vulpis and C. jejuni , highlighting conserved toxin-associated genes despite the lack of confirmed pathogenicity in C. vulpis . The characterization of outer membrane components emphasizes their role in host interactions and immune invasion. Furthermore, evidence of genome plasticity, including HGT, indicates a dynamic evolutionary process driving diversity and pathogenicity. The evolution of pathogenic genes among Campylobacter species may occur through horizontal gene transfer. A number of genomic islands have been identified in Campylobacter species, and key virulence genes such as ( Cdt ) been mapped onto these islands. Viral signatures such as integrated prophages and CRISPR elements were abundant in the genomes of Campylobacter spp., highlighting the high frequency of phage attack and therefore of HGT. Resistome analysis revealed the widespread presence of antibiotic resistance genes, underscoring the clinical significance of Campylobacter as an emerging multidrug-resistant pathogen. Overall, these findings provide an integrative framework that not only enhances our understanding of Campylobacter evolution and pathogenicity, but also providing valuable insights to strengthen future surveillance and therapeutic interventions. Furthermore, future investigations integrating transcriptomic, proteomic, and host-pathogen interaction analyses will be crucial for validating these genomic predictions and identifying novel determinants of pathogenicity and resistance. Future research should also include molecular docking of the Cdt protein to facilitate rational drug design against cytolethal distending toxin. Declarations Competing interests The authors have no relevant financial or non financial interests to disclose. Funding NA Author Contribution This study was conceived and designed by CT. TY performed all the analyses. The manuscript was written by TY and CT. 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FEMS Microbiol Rev. 33(2):376–393. https://doi.org/10.1111/j.1574-6976.2008.00136.x Additional Declarations No competing interests reported. Supplementary Files Supplementarytables.docx Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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13:37:02","extension":"png","order_by":21,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":81101,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage7.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/6d2cdc20d34b4ee4e7a3479d.png"},{"id":94376983,"identity":"24160728-5a09-4263-b9ee-081f467a298c","added_by":"auto","created_at":"2025-10-27 13:36:29","extension":"xml","order_by":22,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":375984,"visible":true,"origin":"","legend":"","description":"","filename":"817ae8ae65ac4ba3a5440986d47527101structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/a63d122e162c24d5c2003c77.xml"},{"id":94376622,"identity":"376431c1-7895-499d-b3d0-39890ae759c0","added_by":"auto","created_at":"2025-10-27 13:35:57","extension":"html","order_by":23,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":396243,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/6bdae22b22a020be53c51153.html"},{"id":94377198,"identity":"123cd025-8350-4b5d-a021-d2fd01f7cdf0","added_by":"auto","created_at":"2025-10-27 13:36:48","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1837326,"visible":true,"origin":"","legend":"\u003cp\u003eAssessment of relationships among \u003cem\u003eCampylobacter\u003c/em\u003espp. based on phylogenetic and phylogenomic methods. (a) Phylogenetic analysis based on 16s rRNA gene sequences using the maximum likelihood method with 1000 bootstrap values. (b) Phylogenetic tree of \u003cem\u003eCampylobacter \u003c/em\u003espp. based on 31 marker proteins formed using the Neighbor-joining method. \u0026nbsp;In (a-b), light pink shades delineate clusters of spp. with major pathogenic species i.e., \u003cem\u003eC. jejuni\u003c/em\u003e, \u003cem\u003eC. coli\u003c/em\u003e and \u003cem\u003eC. fetus\u003c/em\u003e. Phylogenomic dendrogram showing hierarchical clustering of \u003cem\u003eCampylobacter\u003c/em\u003e spp. using (c) whole genome distance-based matrix based on ANI scores (d) Tetra nucleotide frequency. The color of the scale from red to black represents the increasing genome distance on the basis of the two-way ANI matrix. Red denotes the minimum distance and black represents the maximum distance. Species were grouped together into clades on the basis of minimum distance shaded by light pink in ANI and tetra analysis\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/7f8f8b493e3553a30b330769.png"},{"id":94377608,"identity":"04823d33-2bcd-4199-a6e9-b27c80831ca2","added_by":"auto","created_at":"2025-10-27 13:37:21","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1237122,"visible":true,"origin":"","legend":"\u003cp\u003eThe heat map illustrates the comparative functional abundance of metabolic pathways among \u003cem\u003eCampylobacter\u003c/em\u003e species. The clustering of species with \u003cem\u003eC. jejuni\u003c/em\u003e, based on their metabolic pathways, is emphasized, including \u003cem\u003eC. coli, C. hepaticus\u003c/em\u003e, and \u003cem\u003eC. bilis\u003c/em\u003e. The threshold of the number of genes associated with specific pathways is represented through a color scale\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/20ae99e2cd3c47aa960f1a69.jpg"},{"id":94377178,"identity":"6d65ec0c-bbbe-4aaf-a2c3-02dabf22ffc2","added_by":"auto","created_at":"2025-10-27 13:36:44","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":932411,"visible":true,"origin":"","legend":"\u003cp\u003eThe heat map depiction illustrates genes that impart virulence in \u003cem\u003eCampylobacter\u003c/em\u003e spp. The clustering represents interspecies variation in virulence gene repertoires. Major pathogenic species, along with those clustered with them based on the presence of genes associated with virulence factors, are highlighted in red letters. This includes \u003cem\u003eC. vicugnae\u003c/em\u003e, which clusters with \u003cem\u003eC. jejuni\u003c/em\u003e, and \u003cem\u003eC. rectus\u003c/em\u003e, which clusters with \u003cem\u003eC. coli\u003c/em\u003e; however, \u003cem\u003eC. fetus\u003c/em\u003e does not cluster with any species\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/885bed6ac3b1721e7333fa3f.png"},{"id":94376544,"identity":"0979acd6-3c91-4cd0-b9e3-bee2416fefa7","added_by":"auto","created_at":"2025-10-27 13:35:51","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":288887,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eThe graph displays the percentage of identity between the Cdt genes of twenty species belonging to the Campylobacter genus and the Cdt gene of C. jejuni. The bar depicts the identity percentage of Cdt genes of different species with the Cdt gene of C. jejuni with each species denoted by the different colors of the bar. C. vulpis, which is denoted by a blue bar, is most identical to C. jejuni based on Cdt gene similarity\u003c/em\u003e\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/3045f35edc1d6c22970098ce.png"},{"id":94376940,"identity":"c202c058-1e19-4ecf-917d-694140ef8ee8","added_by":"auto","created_at":"2025-10-27 13:36:25","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":13192873,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eThe multiple sequence alignment of the amino acid sequences of subunit B of Cdt genes from various Campylobacter species is presented. A color letters depicts the conserved residues of amino acids among the species of Campylobacter genus. H152, and D185 conserved residues in all species were highlighted. However, D222 conserved in all species except C. bilis\u003c/em\u003e\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/fed14efb6ba42f8e3b52d49b.png"},{"id":94376959,"identity":"35229deb-d1c2-4c48-9a44-c512b460317e","added_by":"auto","created_at":"2025-10-27 13:36:29","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":258701,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003ePhylogenetic assessment of Campylobacter species on the basis of protein sequences of the Cdt gene (in which all three subunits of Cdt gene were present). Red color denotes the major pathogenic species C. jejuni. \u003c/em\u003eThis grouping indicates the \u003cem\u003econservation of toxin related determinants\u003c/em\u003e\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/e961d88c43234e98a9829036.png"},{"id":94377358,"identity":"11e471c7-a648-4e95-876e-db64caadb8d4","added_by":"auto","created_at":"2025-10-27 13:36:59","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":1397140,"visible":true,"origin":"","legend":"\u003cp\u003eType IV secretion system proteins identified in the species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus. The presence of a protein is indicated by the green color, while the absence of a protein is represented by the yellow color\u003c/p\u003e","description":"","filename":"7.png","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/b878ff617a062feb54d5414f.png"},{"id":98428500,"identity":"58f03889-8b89-4021-873f-e5caf6d1503c","added_by":"auto","created_at":"2025-12-17 16:42:05","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":29404493,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/f4be06c6-09e0-4c63-9005-11261919b571.pdf"},{"id":94377107,"identity":"63dc6539-299a-4d00-8938-5b8001808a86","added_by":"auto","created_at":"2025-10-27 13:36:37","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":29933,"visible":true,"origin":"","legend":"","description":"","filename":"Supplementarytables.docx","url":"https://assets-eu.researchsquare.com/files/rs-7771926/v1/c10c6b4cbfc11f4adfbc48fa.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Comparative genomics of the Campylobacter genus: Insights into phylogenomics, virulence, genome plasticity and resistome profiling","fulltext":[{"header":"Introduction","content":"\u003cp\u003e\u003cem\u003eCampylobacter\u003c/em\u003e is a genus of gram-negative bacteria belonging to the phylum Proteobacteria (Marroki et al. 2019). \u003cem\u003eCampylobacter\u003c/em\u003e is slender, curved, spiral, motile, and non-spore-forming microbe (Igwaran et al. 2019). Multiflagella are present in some species such as \u003cem\u003eC. showae\u003c/em\u003e, while in \u003cem\u003eC. gracillis\u003c/em\u003e, flagella are absent (Acke \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Its shape is variable, ranging from 0.5 to 0.9 \u0026micro;m in length (Galanis \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The gastrointestinal tracts of different groups of animals are colonized by species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus (Gendy et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Many species of the genus are associated with animals and humans and most commonly cause a diarrheal infection (Igwaran et al. 2019; Li 2018) known as campylobacteriosis. The most common cause of campylobacteriosis is consumption of contaminated poultry, although humans can also contract \u003cem\u003eCampylobacter\u003c/em\u003e infections from ruminants including cattle, sheep, and goats (Kaakoush et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Chlebicz and Śliżewska, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). \u003cem\u003eCampylobacter\u003c/em\u003e is one of the most prevalent causative agents of food-borne infections in both developed and developing nations (Liu et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Publications from 2014 to 2022 and the official public health websites of 195 nations declared that, Czech Republic had the highest prevalence of campylobacteriosis globally (215 per 100,000 in 2019, followed by Australia (146.8 per 100,000 in 2016) and New Zealand ( 126.1 per 100,000 in 2019) (Liu et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). According to a recent study conducted in Vellore, South India, utilizing PCR analysis on 400 human dysenteric stool samples collected between 2019 and 2020, \u003cem\u003eCampylobacter\u003c/em\u003e was found to be the second most prevalent bacterial enteric pathogen (12%) (Lakshmi et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Additionally, through the use of genus-specific PCR, \u003cem\u003eCampylobacter\u003c/em\u003e spp., were detected in 16.77% of the samples collected from Periurban Bhubaneswar (Mohakhud et el. 2019). Another study performed on children under five in Northeast India between 2014 and 2015 found the prevalence of \u003cem\u003eCampylobacter\u003c/em\u003e infection (10.1%), specifically \u003cem\u003eC. jejuni\u003c/em\u003e (8.1%) in higher abundance (Borkakoty et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In a very small number of cases (1 in 1000) \u003cem\u003eC. jejuni\u003c/em\u003e also causes Gullian-Barre syndrome, an autoimmune neurological disease triggered by the molecular mimicry between outer membrane lipooligosaccharides and human peripheral nerve gangliosides (Li et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Additionally, \u003cem\u003eC. upsaliensis\u003c/em\u003e species of \u003cem\u003eCampylobacter\u003c/em\u003e genus has been isolated from patients with bacteremia, hemolytic-uremic syndrome, spontaneous abortion, and Guillain-Barr\u0026eacute; syndrome (Wang et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe genus \u003cem\u003eCampylobacter\u003c/em\u003e has been divided into major and minor groups of pathogenic species (Rollins et al. 2006), with the major pathogenic species including \u003cem\u003eC. jejuni, C. coli\u003c/em\u003e, and \u003cem\u003eC. fetus\u003c/em\u003e and minor pathogenic species including \u003cem\u003eC. concisus, C. lari, C. upsaliensis\u003c/em\u003e, \u003cem\u003eC. hyointestinalis\u003c/em\u003e and \u003cem\u003eC. hepaticus\u003c/em\u003e (Ienes et al. 2023). Chronic infections with \u003cem\u003eCampylobacter\u003c/em\u003e spp. have also been observed to sometimes lead to colorectal cancer, Barrett's esophagus, and mucosa associated lymphoid tissue lymphoma (Marroki et al. 2019; Kato et al. 2021). Apart from the major and minor pathogenic species, some other members of the genus such as \u003cem\u003eC. ureolyticus\u003c/em\u003e (Bullman et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Bullman et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2012\u003c/span\u003e), C. \u003cem\u003ehelveticus, C. insulanigrae, C. mucosalis\u003c/em\u003e (Inglis et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), C. \u003cem\u003erectus\u003c/em\u003e (Lastovica et al. 2024) and \u003cem\u003eC. sputorum\u003c/em\u003e have been associated with gastroenteritis (Lindblom et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Man et al. 2011). Moreover, \u003cem\u003eC. fetus\u003c/em\u003e has been shown to cause extra-intestinal infections and septic abortion in farm animals (Allos et al. 1995; Wu et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) Additionally, \u003cem\u003eC. showae\u003c/em\u003e associated with crohns disease and ulcerative colitis (Man et al. 2011). A comparative genomic analysis of thirty-nine \u003cem\u003eCampylobacter\u003c/em\u003e species has been conducted to date. This analysis identified antibiotic resistance genes, virulence factor genes, and pangenome genes. It also examined genetic diversity, evolutionary traits, and evaluated genome plasticity.\u003c/p\u003e\u003cp\u003eAmong the pathogenic species of \u003cem\u003eCampylobacter, C. jejuni\u003c/em\u003e contributes for almost 90% of the instances of campylobacteriosis, while \u003cem\u003eC. coli\u003c/em\u003e is responsible for less than ten percent of cases (Heimesaat et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Liu et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Numerous studies have linked prolonged gastrointestinal infections caused by \u003cem\u003eC. jejuni\u003c/em\u003e to the development of colorectal cancer (Blaser et al. 2008; Guerra et al. 20011; He et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Kato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Duijster et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and mucosa associated lymphoid tissue (MALT) lymphoma (Louwen et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). \u003cem\u003eC. jejuni\u003c/em\u003e produces a genotoxic substance known as cytolethal distending toxin (CDT), having DNAse activity (Lai et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; He et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; and Chen et al. 2024). CDT activity causes double-stranded DNA breaks and increases the probability of transformation of normal cells into cancer cells (Brauner et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Vogtmann et al. 2016; Tremblay et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Apart from CDT, various other virulence factors are involved in the establishment and continuation of infection by \u003cem\u003eC. jejuni\u003c/em\u003e (Kemper et al. 2023). These virulence factors are related with bacterial adherence to intestinal mucosa, flagella mediated motility, invasive capability, chemotaxis, quorum sensing and outer membrane component lipopolysacchaides (LPS).\u003c/p\u003e\u003cp\u003eEven though a number of species of this genus have been identified as pathogenic (major or minor), the pathogenic potential of most of the species has not yet been investigated. This study was designed with the aim of elucidating the phylogenomic relationships between different species of the genus \u003cem\u003eCampylobacter\u003c/em\u003e and the analysis of their pathogenic potential by the comparative analysis of their virulence genes, antibiotic resistance genes and metabolic pathways. Apart from this, we have identified the genomic islands; prophage regions and CRISPRs analysis in order to determine the genomic plasticity among the species of \u003cem\u003eCampylobacter\u003c/em\u003e genus.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003ePhylogenomic analysis\u003c/h2\u003e\u003cp\u003eTo perform phylogenomic analysis, the genomes of fifty \u003cem\u003eCampylobacter\u003c/em\u003e species whose genome sequences were publicly available were downloaded from the NCBI database (Table \u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Phylogenetic analysis based on conventional 16s rRNA gene sequences was performed, for which the RNAmmer version 1.2 server was used to extract 16s rRNA genes from each genome (Lagesen et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). A phylogenetic tree was constructed using the maximum likelihood algorithm (Felsenstein et al. 1993) with a 1000 bootstrap value in MEGA X. To assure the accuracy of the phylogenetic relationship multiple conserved bacterial marker protein based phylogenetic analysis was performed, for which 31 conserved bacterial marker proteins were extracted from each of the 50 species of \u003cem\u003eCampylobacter\u003c/em\u003e using AmphoraNet server (Kerepesi et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). A phylogenetic tree was constructed using the Neighbor Joining algorithm (Saitou et al.1987) with 1000 bootstrap revaluations implemented in MEGA X (Erickson et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Further, whole genome data-based phylogenomic analysis was performed, which includes Average nucleotide identity (ANI) and tetranucleotide frequency. ANI values were calculated by employing the BLASTALL algorithm of J species server version 1.2.1. (Richter et al. 2009). To calculate ANI values and tetra nucleotide frequency, genomes from each species were uploaded to J species server version 1.2.1. After that two-way pair matrix was formed containing pairwise ANI scores. Hierarchial clustering was performed and dendogram was constructed using MeV 2.0 software for ANI and tetranucleotides frequency scores.\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\u003eAccession numbers, isolation sources and general genomic features of \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"10\"\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=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies name\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eStrain\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNCBI accession number\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSource of isolation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eGenome size (in Mb\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eGC Content\u003c/p\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNumber of contings\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePlasmid\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003etRNAs\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003eNumber of coding sequences\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCampylobacter jejuni\u003c/em\u003e\u003c/p\u003e\u003cp\u003eGundogdu et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2007\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC 70089\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNC_002163.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNot known\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e30.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1687\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. armoricus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG; Yee 2020\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCCUG 73571\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053825.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eRiver water\u003c/p\u003e\u003cp\u003eFrance: Brittany\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1625\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. avium\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller 2017(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG\u003c/p\u003e\u003cp\u003e24591\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP022347.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCaecal\u003c/p\u003e\u003cp\u003ehost-chicken\u003c/p\u003e\u003cp\u003eItaly: Bolognan\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1783\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. concisus\u003c/em\u003e Miller WG 2016 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e33237\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP012541.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eGingival sulcus\u003c/p\u003e\u003cp\u003ehost-homo sapiens\u003c/p\u003e\u003cp\u003eUSA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e37.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1898\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. coragiensis\u003c/em\u003e Miller WG 2020\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG27932\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053842.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003ehost-Macaca silenus\u003c/p\u003e\u003cp\u003eIreland: Cork\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e31.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e38 Kb (NZ_CP053843.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1767\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. coli\u003c/em\u003e\u003c/p\u003e\u003cp\u003eKane et al. 2017(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAerotolerant OR12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP019977.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOrganic chicken farm United Kingdom: Lincolnshire\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e30.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2196\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. curvus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e35224\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053826.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHuman, jaw abscess\u003c/p\u003e\u003cp\u003eUSA: Massachusetts\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e44.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2010\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hominis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eFouts 2007 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC BAA-381\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNC_009714.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNot known\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e31.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e36 Kb (NC_009713.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1738\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. fetus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eDavis et al. 2021\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCF00A031\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP059443.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePreputial wash\u003c/p\u003e\u003cp\u003eHost \u0026ndash;bovine\u003c/p\u003e\u003cp\u003eCanada:British Columbia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e26 Kb (NZ_CP059444.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1811\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. geochelonis\u003c/em\u003e\u003c/p\u003e\u003cp\u003e(direct submission 2016)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRC7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_FIZQ01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCloacal swab\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2070\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. gracilis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller et al. 2015\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e33236\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP012196.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOral\u003c/p\u003e\u003cp\u003eHost-Homo sapiens\u003c/p\u003e\u003cp\u003eUSA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e46.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2757\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. helveticus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2017 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e51209\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP020478.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost-cat\u003c/p\u003e\u003cp\u003eSwitzerland-Berne\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1. 72 Kb (NZ_CP020479.1)\u003c/p\u003e\u003cp\u003e2. 34 Kb(NZ_CP020480.1)\u003c/p\u003e\u003cp\u003e3. 32 Kb\u003c/p\u003e\u003cp\u003e(NZ_CP020481.1)\u003c/p\u003e\u003cp\u003e4. 16 Kb(NZ_CP020482.1)\u003c/p\u003e\u003cp\u003e5. 17 Kb\u003c/p\u003e\u003cp\u003e( NZ_CP020483.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2028\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hepaticus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eVan et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2019\u003c/span\u003e (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eHV10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP031611.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eGallus gallus (whole organism)\u003c/p\u003e\u003cp\u003eAustralia, Victoria\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1519\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hyointestinalis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2020\u003c/p\u003e\u003cp\u003e(Direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLawsoni CHY5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053828.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProcrine stomatch\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e44 Kb (NZ_CP053829.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1880\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. iguaniorum\u003c/em\u003e Gilbert et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2016\u003c/span\u003e (Direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2463D\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP010995.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eIguana iguana\u003c/p\u003e\u003cp\u003eNetherlands\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1895\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. insulaenigrae\u003c/em\u003e Miller et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNTCC\u003c/p\u003e\u003cp\u003e12927\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP007770.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eMarine mammal\u003c/p\u003e\u003cp\u003eUK Scotland\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1485\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. lanienae\u003c/em\u003e Miller et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNCTC\u003c/p\u003e\u003cp\u003e13004\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP015578.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003ehost-homo sapiens\u003c/p\u003e\u003cp\u003eSwitzerland\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1634\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. lari\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRM\u003c/p\u003e\u003cp\u003e2100\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNC_012039.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eNot known\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1. 46 Kb (NC_012040.1)\u003c/p\u003e\u003cp\u003e2. 18 Kb ( CP000933.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. Mucosalis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e43264\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053831.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePig, small intestine\u003c/p\u003e\u003cp\u003eUK\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e36.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1834\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. ornithocola\u003c/em\u003e Miller WG 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG\u003c/p\u003e\u003cp\u003e29815\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053848.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eWild bird fecal samples\u003c/p\u003e\u003cp\u003eChile: Valdivia\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1627\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC.pinnipediorum\u003c/em\u003e Miller WG 2016\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRM172661\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP012547.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSea lion lung\u003c/p\u003e\u003cp\u003eUSA: California\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e30.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1775\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. rectus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2016\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC\u003c/p\u003e\u003cp\u003e33238\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP012543.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOral\u003c/p\u003e\u003cp\u003ehost-homosapiens, USA: Boston\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e44.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e3083\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. showae\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2016\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eATCC 51146\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP012544.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOral\u003c/p\u003e\u003cp\u003ehost-homosapiens, Japan: Showa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e45.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2312\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. novaezeelandiae\u003c/em\u003e Bloomfield; Wilkinson D.A. 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eB423b\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_QPGR01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAnas platyrhynchos,\u003c/p\u003e\u003cp\u003eNew Zealand\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e27.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e59\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e39\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1645\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. sputorum\u003c/em\u003e Miller et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG\u003c/p\u003e\u003cp\u003e11764\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP019684.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHuman, fecal\u003c/p\u003e\u003cp\u003eCanada: Ottawa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1765\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. upsaliensis\u003c/em\u003e Miller 2020\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRM3940\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053849.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eHuman, fecal\u003c/p\u003e\u003cp\u003eUSA: Los Angeles, California\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1668\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. ureolyticus\u003c/em\u003e Kyrpides et al. 2013 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDSM\u003c/p\u003e\u003cp\u003e20703\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_KB894730.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAmniotic fluid,\u003c/p\u003e\u003cp\u003eCanada\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1803\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. volucris\u003c/em\u003e Miller et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2014\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG\u003c/p\u003e\u003cp\u003e24379\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP007774.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eBlack headed gull,\u003c/p\u003e\u003cp\u003eSweden\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1520\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. subantarcticus\u003c/em\u003e Miller et al.2014\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG\u003c/p\u003e\u003cp\u003e24377\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP007773.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eGrey headed albatross,\u003c/p\u003e\u003cp\u003eSouth Georgia and the South Sandwich\u003c/p\u003e\u003cp\u003eIslands\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1918\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. vulpis\u003c/em\u003e\u003c/p\u003e \u003cp\u003eParisi 2015. (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e73/13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_LDWY01000065.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFox blind gut, Italy: Monterenzio\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1667\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. protucalensis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eSilva et al.\u003c/p\u003e\u003cp\u003e2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFMV-PI01\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eVWSJ01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eBovine preputial sample,\u003c/p\u003e\u003cp\u003ePortugal: Lisbon\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e98\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e39\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1798\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCandidates C. infans\u003c/em\u003e\u003c/p\u003e\u003cp\u003eDuim 2021 ( direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e19S00001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP049075.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost-homosapiens, Netherlands: Utrecht\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e5.8 Kb (NZ_CP049076.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1871\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. taeniopygiae\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMannion 2018\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMIT10-5678\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_NXLY01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost- Taeniopygia guttata,\u003c/p\u003e\u003cp\u003eUSA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e88\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1824\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. estrildidarum\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMannion 2018 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMIT17-664\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_NXLZ01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost- Taeniopygia guttata,\u003c/p\u003e\u003cp\u003eUSA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1798\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. aviculae\u003c/em\u003e Mannion 2018 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMIT17-670\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_NXMA01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost- Taeniopygia guttata,\u003c/p\u003e\u003cp\u003eUSA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e59\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1752\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. canadensis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2019\u003c/p\u003e\u003cp\u003e(direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG 24001\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP035946.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ecloacal swab\u003c/p\u003e\u003cp\u003ehost- Whooping crane\u003c/p\u003e\u003cp\u003eCanada: Calgary\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e27.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1885\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. troglodytis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMannion A\u003c/p\u003e\u003cp\u003e2018 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMIT 05-9149A\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_QHLI01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eChimpanzees, USA: MIT Cambridge, MA\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e295\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e4083\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. blaseri\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller WG 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG 30333\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP053841.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFeces\u003c/p\u003e\u003cp\u003eHost- Phoca vitulina,\u003c/p\u003e\u003cp\u003eNetherlands:Pieterburen\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e29.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1887\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. pleoridis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eLane 2020 (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2016D-0074\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP063079.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eShellfish\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e28.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e45 Kb (NZ_CP063080.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1665\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. cuniculorum\u003c/em\u003e Miller et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e\u003c/p\u003e\u003cp\u003e(direct submission).\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDSM 23162\u0026thinsp;=\u0026thinsp;LMG 24588\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP020867.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eRabbit, caecal,\u003c/p\u003e\u003cp\u003eItaly: Bologna\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e31.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1. 49 Kb (NZ_CP020868.1)\u003c/p\u003e\u003cp\u003e2. 18 Kb (NZ_CP020869.1)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1996\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. analoticus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eAydin et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003efaydin-G140\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_JAGSSY010000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eFaeces of Anatolian Ground squirrel in Turkey\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e35.2%\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1968\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. bilis\u003c/em\u003e\u003c/p\u003e\u003cp\u003ePhung et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2022\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eVicNov18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eWUED01000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eChikens with spotty liver disease\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e30.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1568\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. cryaerophila\u003c/em\u003e\u003c/p\u003e\u003cp\u003eNeill et al. \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e1985\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG 24291\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNXGK01000077.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eBovine Brain, aborted fetus, Ireland\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e27.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e91\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2145\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. devanensis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e (direct submission)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eNCTC 13003\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNZ_CP018788.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eUnkown\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1618\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. magnus\u003c/em\u003e\u003c/p\u003e\u003cp\u003eGruntar et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2023\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e46386\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eJAQSLK010000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCaecal contents of domestic pigs.\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e38.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1862\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. majalis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eLynch et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG 7974\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCAJHOF010000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProcelli gastro-intestinal mucosa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e59\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e39\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2086\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. massiliensis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eAntezacker et al. 2021\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eMarseille-Q3452\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eJACLZK010000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eDental source host-homo sapiens, France\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e45.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e2452\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. procelli\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2024\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRM6137\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCP018789.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eWild pig, fecal\u003c/p\u003e\u003cp\u003eUSA: California\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e34.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1674\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. suis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eLynch et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2022\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLMG 8286\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCAJHOE010000001.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eProcrine gastro-intestinal mucosa\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e37.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1827\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. vicugnae\u003c/em\u003e\u003c/p\u003e\u003cp\u003eMiller et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2024\u003c/span\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eRM12175\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eCP018793.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAlapacs goats and sheep\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e32.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e\u003cp\u003e1669\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"10\"\u003eND*-not determined\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eFunctional potential analysis\u003c/h3\u003e\n\u003cp\u003eFunctional flexibility among \u003cem\u003eCampylobacter\u003c/em\u003e spp. was determined by comparative genomic analysis, which is based on metabolic pathways. For this, amino acid sequences of all the fifty species of \u003cem\u003eCampylobacte\u003c/em\u003er were retrieved from the RAST server (Aziz et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), followed by the gene finding using KAAS (KEGG Automatic Annotation Server) by BLAST against the KEGG (Kyoto Encyclopedia of Genes and Genomes) database (Ogata et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). Metabolic pathways were reconstructed using MinPath (Minimal Set of Pathways) (Ye et al. 2009). One-way hierarchical clustering was performed based on 125 metabolic pathways. For hierarchical clustering, a heat map was created using the Euclidian and Ward algorithms, followed by a matrix transformation of all values with Log 10 and keeping 500 of the rows with the highest standard deviation (Ryan et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eAnalysis of the genes associated with virulence factors\u003c/h3\u003e\n\u003cp\u003ePrevious studies have identified the presence of virulence factors in \u003cem\u003eCampylobacter\u003c/em\u003e genus (Igwaran et al. 2019; Kemper et al. 2023; Bunduruș et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). These virulence factors were identified in the genomes of different species of this genus, using the annotation generated by RAST and a heat map was generated by the NGCHM builder (Ryan et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eCytolethal distending toxin (Cdt) is a key virulence factor found in \u003cem\u003eCampylobacter\u003c/em\u003e species with potential link to colorectal cancer (Kato et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). As a result, \u003cem\u003eCdt\u003c/em\u003e genes were examined in each species within the genus. Species containing \u003cem\u003eCdt\u003c/em\u003e genes were subjected to a BLASTn analysis (Altschul et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) against the \u003cem\u003eCdt\u003c/em\u003e genes of \u003cem\u003eC. jejuni\u003c/em\u003e to assess the similarity of \u003cem\u003eCdt\u003c/em\u003e genes from other \u003cem\u003eCampylobacter\u003c/em\u003e species to \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni.\u003c/em\u003e For this analysis subunit A, B and C of each species concatenated and compared. Further, a multiple sequence alignment of Cdt protein of subunit B sequences was conducted using Muscle (see Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e) to identify the conserved domains within the protein. A phylogenetic tree was constructed based on the amino acid sequence (concatenated sequence of subunit A, B and C) of the \u003cem\u003eCdt\u003c/em\u003e gene using the Neighbor-Joining method in Mega X software.\u003c/p\u003e\u003cp\u003eType IV ((T4SS) and Type VI secretion systems(T6SS) play a crucial role in host-pathogen interactions and are present in several well-known bacterial pathogens, including \u003cem\u003eSalmonella, Pseudomonas, Yersinia\u003c/em\u003e, and \u003cem\u003eVibrio\u003c/em\u003e (Pukatzki et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Consequently, proteins associated with Type IV and Type VI secretion systems have been identified in each \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003eThe outer membrane components of \u003cem\u003eCampylobacter\u003c/em\u003e serve as multifunctional virulence factors, playing a crucial role in the bacterium's ability to colonize, invade, and survive within the human host (Lopes et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). To classify lipopolysaccharides (LPS) from \u003cem\u003eCampylobacter\u003c/em\u003e species based on their components, we examined the structures of lipid A, the outer core, and the inner core in each species of the genus. These were categorized into rough LPS, deep rough LPS, and smooth-type LPS. Additionally, sialyated carbohydrate lipooligosaccharide (LOS) structures synthesis enzymes on the outer membranes of the members of this genus were identified as they have been known to be associated with complement-mediated nerve fatalities and severe colitis in \u003cem\u003eC. jejuni\u003c/em\u003e (Louwen et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Further, LOS components identified were compared to the LOS of \u003cem\u003eC. jejuni\u003c/em\u003e to assess their similarity.\u003c/p\u003e\n\u003ch3\u003eGenome plasticity and resistome analysis\u003c/h3\u003e\n\u003cp\u003eGenomic islands, along with other mobile and accessory genetic elements, play a significant role in the horizontal movement of some genes in bacterial populations and contribute to the genomic flexibility of bacteria (Dobrindt et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Genomic islands were identified using Genomic Island Viewer 4 (Bertelli et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) based on variations in GC content and codon usage bias and IspandPath-DIMOB. The presence of prophage regions within bacterial genomes can signify the occurrence of horizontal gene transfer in bacteria (Borodovich et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The detection of prophage regions in \u003cem\u003eCampylobacter\u003c/em\u003e species was conducted using the PHASTER server (Arndt et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Subsequently, to ascertain which genes in \u003cem\u003eCampylobacter\u003c/em\u003e species were acquired through phages, bacterial gene sequences integrated with prophage regions were identified. Furthermore, CRISPR genes were extracted from each genome by uploading the genomes to the CRISPRFinder online server (Grissa et al. \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). This server conducted a BLAST search against the dbCRISPR database, categorizing the results into true and false CRISPRs based on their association with CRISPR-associated (cas) genes. Additionally, the spacer sequences of \u003cem\u003eCampylobacter\u003c/em\u003e species were subjected to a BLAST search against the NCBI virus database to identify similarities with viral sequences, thereby predicting which viruses may be involved in horizontal gene transfer within \u003cem\u003eCampylobacter\u003c/em\u003e species. Furthermore, antibiotic resistance genes were identified in the fifty species of \u003cem\u003eCampylobacter\u003c/em\u003e genus using the resistance gene identifier 6.0.1 (Kwatra \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e\u003ch2\u003ePhylogenomic analysis\u003c/h2\u003e\u003cp\u003eA general analysis of the genomic characteristics of 50 \u003cem\u003eCampylobacter\u003c/em\u003e species indicated that their genome sizes ranged from 1.4 Mb to 2.9 Mb, with GC content between 27.2% and 46.5%. The species and genomes examined in this study were predominantly isolated from food sources, including meat, poultry, and animal-associated environments. Among the 50 species of the genus, plasmids were identified in nine species, with plasmid sizes ranging from 16 Kb to 58 Kb (Table\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Plasmid encoded genes associated with type IV secretion system in \u003cem\u003eC. fetus, C. pleoridis, C. helveticus\u003c/em\u003e and \u003cem\u003eC. lari\u003c/em\u003e species. Phylogenetic analysis based on the 16S rRNA genes revealed that \u003cem\u003eC. jejuni\u003c/em\u003e and \u003cem\u003eC. coli\u003c/em\u003e, which are major pathogenic species, form a monophyletic clade. In addition, \u003cem\u003eC. fetus\u003c/em\u003e (major pathogenic species) and \u003cem\u003eC. iguaniorum\u003c/em\u003e (non-pathogenic species) formed a monophyletic clade. However, \u003cem\u003eC. cryaerophila\u003c/em\u003e is distinct among 50 spp. of the \u003cem\u003eCampylobacter\u003c/em\u003e. To further elucidate the phylogenetic relationships among species within the genus, phylogenetic analyses were conducted using 31 conserved bacterial marker genes. These analyses revealed that \u003cem\u003eC. coli, C. hepaticus\u003c/em\u003e, and \u003cem\u003eC. jejuni\u003c/em\u003e spp. are the most closely related species, forming a monophyletic clade. Additionally, other species, including \u003cem\u003eC. helveticus, C. upsaliensis\u003c/em\u003e (minor pathogenic species), and \u003cem\u003eC. vulpis\u003c/em\u003e, are closely related and grouped within a single clade, while \u003cem\u003eC. fetus, C. iguaniorum\u003c/em\u003e, and \u003cem\u003eC. hyointestinalis\u003c/em\u003e constitute another monophyletic clade. ANI provides strong resolution for genomes that share 80\u0026ndash;100% ANI, indicating they belong to the same species, as well as for closely related species that share less than 80% ANI. In the present study, the ANI values ranged from 68 to 72% among \u003cem\u003eCampylobacter\u003c/em\u003e spp. Whole-genome analyses utilizing ANI and TETRA frequency metrics have demonstrated that \u003cem\u003eC. jejuni, C. coli, C. hepaticus\u003c/em\u003e, and \u003cem\u003eC. taniopygae\u003c/em\u003e are species with close phylogenetic relationships (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cb\u003eComparative analysis of functional repertoire of\u003c/b\u003e \u003cb\u003eCampylobacter\u003c/b\u003e \u003cb\u003egenus\u003c/b\u003e\u003c/p\u003e\u003cp\u003eReconstruction of metabolic pathways for the 50 species within the \u003cem\u003eCampylobacter\u003c/em\u003e genus, based on KAAS, led to the identification of several conserved metabolic pathways, including glycolysis, the tricarboxylic acid (TCA) cycle, the pentose phosphate pathway, pentose and glucuronate interconversion, fructose and mannose metabolism, and the galactose metabolism pathway. Notably, the fatty acid degradation pathway was absent in \u003cem\u003eCandidatus C. infans, C. laninae\u003c/em\u003e, and \u003cem\u003eC. avium\u003c/em\u003e. Additionally, certain pathways were absent in specific species; for instance, the histidine metabolism pathway was present in all species except \u003cem\u003eC. taeniopygae\u003c/em\u003e. The benzoate degradation pathway was absent in \u003cem\u003eC. curvus\u003c/em\u003e but present in all other species. Furthermore, the beta-alanine metabolic pathway was not found in \u003cem\u003eC. taeniopygae, C. aviculae\u003c/em\u003e, and \u003cem\u003eC. canadensis\u003c/em\u003e. With the exception of \u003cem\u003eC. hominis\u003c/em\u003e, all \u003cem\u003eCampylobacter\u003c/em\u003e species exhibited pathways for dioxin and xylene degradation, biofilm formation (\u003cem\u003ePseudomonas aeruginosa\u003c/em\u003e), and unsaturated fatty acid biosynthesis. Moreover, the glucose inotate biosynthesis pathway was absent in \u003cem\u003eC. novazeelandiae\u003c/em\u003e and \u003cem\u003eC. taeniopygae\u003c/em\u003e. Similarly, the mineral absorption pathway was exhibited by all species but was absent in \u003cem\u003eC. blaseri\u003c/em\u003e. Ascorbate and validomycin biosynthesis pathways were present in \u003cem\u003eC. coli, C. geochelonsis, C. helveticus, C. insulanigare\u003c/em\u003e, and \u003cem\u003eC. troglodytis\u003c/em\u003e, but absent in other species. Some metabolic pathways associated with the pathogenicity of the bacterium and antibiotic resistance were found to be present in all species of the genus, including the bacterial secretion system, bacterial chemotaxis, quorum sensing, two-component system, lipopolysaccharide synthesis, and biofilm formation \u003cem\u003eE. coli\u003c/em\u003e, platinum drug resistance, antifolate resistance, cationic antimicrobial peptide resistance vanomycin resistance and beta-lactum resistance. Hierarchical clustering of the metabolic pathways across the 50 genomes revealed a close relationship among \u003cem\u003eC. jejuni, C. coli, C. hepaticus\u003c/em\u003e, and \u003cem\u003eC. bilis\u003c/em\u003e, as corroborated by phylogenomic analysis (Fig.\u0026nbsp;2).\u003c/p\u003e\u003cp\u003e\u003cb\u003eFigure\u0026nbsp;2\u003c/b\u003e The heat map illustrates the comparative functional abundance of metabolic pathways among \u003cem\u003eCampylobacter\u003c/em\u003e species. The clustering of species with \u003cem\u003eC. jejuni\u003c/em\u003e, based on their metabolic pathways, is emphasized, including \u003cem\u003eC. coli, C. hepaticus\u003c/em\u003e, and \u003cem\u003eC. bilis\u003c/em\u003e. The threshold of the number of genes associated with specific pathways is represented through a color scale\u003c/p\u003e\u003c/div\u003e\n\u003ch3\u003eComparative analysis of factors associated with virulence\u003c/h3\u003e\n\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\u003ch2\u003e(I) Genes associated with virulence\u003c/h2\u003e\u003cp\u003eVirulence factors impart specific traits associated with virulence, including bacterial adherence to the intestinal mucosa, motility mediated by flagella, and the ability to invade host tissues. Genes involved in imparting virulence (n\u0026thinsp;=\u0026thinsp;37) that were identified in \u003cem\u003eCampylobacter\u003c/em\u003e genus included chemotaxis virulence factors CheA, CheB, CheR, CheV, CheW, CheZ Cet A, Cet B, LuxS, motility virulence factors JlpA, flaB, flaA, Flagellin C, major outer membrane protein, outer membrane protein CadF, adhesion protein A CapA, periplasmic binding protein PEB1A, fibronectin-like protein A FlpA, type IV secretion system, type VI secretion system, fibronectin F, racR, dnaJ, docA, flagellin C, \u003cem\u003eCampyloabcter\u003c/em\u003e invasion antigen B CiaB, invasion antigen C, \u003cem\u003eCampylobacter\u003c/em\u003e invasion antigen CiaD, invasion associated protein iamA, periplasmic protein HtrA, expression of invasion pldA, sialyltransferase, intracellular invasion Cial, Lipopolysaacharide production WlaN, Sialyltransferase, methyl accepting chemotaxis protein, and cytolethal distending toxin. Species clustering on the basis of these factors demonstrated the close associatoion of \u003cem\u003eC. jejuni\u003c/em\u003e, with \u003cem\u003eC. vicugnae\u003c/em\u003e and \u003cem\u003eC. coli\u003c/em\u003e with \u003cem\u003eC. rectus\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Among the major pathogenic species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus, twenty nine, twenty six and twenty four virulence factors have been identified in \u003cem\u003eC. jejuni, C. coli\u003c/em\u003e and \u003cem\u003eC. fetus\u003c/em\u003e respectively. Additionally, among the minor pathogenic species, nineteen, twenty two, twenty five, twenty six and twenty virulence factors were identified in \u003cem\u003eC. concisus\u003c/em\u003e, \u003cem\u003eC. hyointestinalis\u003c/em\u003e, \u003cem\u003eC. lari\u003c/em\u003e, \u003cem\u003eC. upsaliensis\u003c/em\u003e and \u003cem\u003eC. hepaticus\u003c/em\u003e respectively. High number of virulence factor genes were also identified in species of the genus, which have not been classified as major or minor pathogens but have been reported to be associated with gastroenteritis, including \u003cem\u003eC. heleveticus\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;29), \u003cem\u003eC. insulanigare\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;24) and \u003cem\u003eC. mucosalis\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;26), \u003cem\u003eC. sputorum\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;22), \u003cem\u003eC. rectus\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;20), \u003cem\u003eC. ureolyticus\u003c/em\u003e (n\u0026thinsp;=\u0026thinsp;10) and \u003cem\u003eC. showae\u003c/em\u003e (=\u0026thinsp;22) have been associated with colitis. Furthermore, high number of virulence factor genes (n\u0026thinsp;\u0026ge;\u0026thinsp;19) were also identified in species which have not been reported as pathogenic till date including \u003cem\u003eC. armoricus, C. avium, C. iguaniorum, C. laninae, C. ornithocola, C. pinnipediorum, C. novaezeelandiae, C. subantracticus, C. volucris, C. taeniopygae, C. estrildidarum, C. troglodytis, C. vulpis, C. aviculae, C. pleridids\u003c/em\u003e, \u003cem\u003eC. cuniculorum\u003c/em\u003e, and \u003cem\u003eC. procelli\u003c/em\u003e and \u003cem\u003eC. vicugnae.\u003c/em\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003ePrevious studies have indicated the association of cytolethal distending toxin (\u003cem\u003eCdt\u003c/em\u003e) of \u003cem\u003eCampylobacter jejuni\u003c/em\u003e with initiation of colon cancer in patients suffering from chronic infections (He et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; He et al. \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Guidi. 2014; Graillot et al. \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Hartl et al. 2020; Yusuf et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In this study, we identified and compared the \u003cem\u003eCdt\u003c/em\u003e genes across 50 \u003cem\u003eCampylobacter\u003c/em\u003e species. This analysis revealed that 27 out of the 50 species harbor \u003cem\u003eCdt\u003c/em\u003e genes, specifically \u003cem\u003eC. jejuni, C. coli\u003c/em\u003e, and \u003cem\u003eC. fetus\u003c/em\u003e (major pathogens); \u003cem\u003eC. lari, C. upsaliensis\u003c/em\u003e and \u003cem\u003eC. hyointestinalis\u003c/em\u003e (minor pathogens); \u003cem\u003eC. helveticus, C. insulanigrae\u003c/em\u003e and \u003cem\u003eC. mucosalis\u003c/em\u003e (associated with gastroenteritis); \u003cem\u003eC. vulpis, C. taeniopygae, C. armoricus, C. aviculae, C. estrildidarum, C. subantracticus, C. pleoridis, C. volucris, C. geochelonsis, C. iguaniorum, C. laninae,, C. devanensis, C. procelli, C avium, C. ornithocola, C. canadensis, C. vicugnae\u003c/em\u003e and \u003cem\u003eC. bilis\u003c/em\u003e (pathogenicity has not been reported yet). Cdt consists of three subunits: A, B and C, with subunit B being associated with DNase activity (Brauner et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). The gene encoding for subunit B of the \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni\u003c/em\u003e is composed of 798 bp in size, the gene for subunit A consists of 807 bp, and that of subunit C is made up of 570 bp. Genes encoding for all three subunits were identified in 22 species. The gene for subunit C was not identified in \u003cem\u003eC. iguaniorum\u003c/em\u003e, \u003cem\u003eC. canadensis\u003c/em\u003e, and that for subunit A was not located in \u003cem\u003eC. avium, C. vicugnae\u003c/em\u003e and \u003cem\u003eC. devanensis\u003c/em\u003e indicating non-functional Cdt proteins in these species. The BLASTn analysis of \u003cem\u003eCdt\u003c/em\u003e nucleotide sequences of 22 \u003cem\u003eCampylobacter\u003c/em\u003e species (in which all three subunits were present) against the \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni\u003c/em\u003e revealed that the \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. bilis\u003c/em\u003e exhibited highest similarity with \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni\u003c/em\u003e with 74.05% identity e-value 2e-75) however, its nucleotide sequences of B subunit was truncated (size of subunit B of \u003cem\u003eC. bilis\u003c/em\u003e was 573 bp). Furthermore, \u003cem\u003eC. vulpis\u003c/em\u003e exhibited the highest similarity to the \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni\u003c/em\u003e (71.70% identity; e-value 0.0) followed by \u003cem\u003eC. helveticus\u003c/em\u003e (71.34% identity; e-value 0.0) and \u003cem\u003eC. upsaliensis\u003c/em\u003e (70.78% identity; e-value 0.0). The identity percentage of the \u003cem\u003eCdt\u003c/em\u003e genes of other species with that of \u003cem\u003eC. jejuni\u003c/em\u003e ranged from 69.53% to 64.57% (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Furthermore, multiple sequence alignment of the amino acid sequences of \u003cem\u003eCdt\u003c/em\u003e genes from \u003cem\u003eCampylobacter\u003c/em\u003e species revealed the conserved regions on catalytic sites of subunit B that may be critical for the function of the Cdt protein across different species including, H152, D185 and D222 ( D222 conserved in all spp. except \u003cem\u003eC. bilis\u003c/em\u003e) (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e5\u003c/span\u003e). The phylogenetic tree based on protein sequences of the \u003cem\u003eCdt\u003c/em\u003e gene demonstrated the \u003cem\u003eC. jejuni\u003c/em\u003e formed a distinct cluster, separated from the other \u003cem\u003eCampylobacter\u003c/em\u003e species. The distinct clustering pattern indicates that \u003cem\u003eCdt\u003c/em\u003e gene in \u003cem\u003eC. jejuni\u003c/em\u003e has undergone unique evolutionary pressure, which may contribute to its well- established pathogenic potential compared to other members of the genus (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e6\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eBoth T4SS and T6SS play important roles in pathogenicity (Costa et al. 2019). Analysis of T4SS proteins in each species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus revealed that T4SS proteins were present in 29 out of the 50 species. However, not all species included the complete set of system proteins (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e7\u003c/span\u003e). Examination of the T6SS genes in \u003cem\u003eCampylobacter\u003c/em\u003e species led to their identification in 21 out of the 50 species (supplementary table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003e1\u003c/span\u003e online resource). Generally; 13\u0026ndash;15 core genes form the cluster encoding T6SS.\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e\u003ch2\u003e(II) Outer membrane pathogenic component analysis\u003c/h2\u003e\u003cp\u003eThe lipopolysaccharides that were identified among the species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus were classified based on their components into rough LPS (lipid A and outer core), deep rough LPS (lipid A and inner core), and smooth LPS (complete LPS: lipid A, outer core, inner core, and O-antigen). Among the species of the genus, 41 species have deep rough LPS, making them highly susceptible to hydrophobic antibiotics. \u003cem\u003eC. jejuni, C. armoricus, C. laninae, C. hyointestinalis, C. upsaliensis, C. bilis, C. cryaerophila, C. devanensis\u003c/em\u003e and \u003cem\u003eC. vicugnae\u003c/em\u003e were found to have rough LPS, making it less susceptible to antibiotics (Table \u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Furthermore, sialylated oligosaccharide structures synthesis enzymes including ADP-heptose-lipooligosaccharide heptosyl transferase, CMP-Neu5Ac synthase and CMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase were examined in each species of the \u003cem\u003eCampylobacter\u003c/em\u003e genus to assess their potential for pathogenicity based on their membrane components. Forty one species were found to harbour the sialylated lipooligosaccharide synthesis enzyme known as ADP-heptose-lipooligosaccharide heptosyl transferase. This finding suggests that presence of sialyated oligosaccharide structure in these forty one species may have incorporated in their virulence. Additionally, the BLAST result of ADP-heptose-lipooligosaccharide heptosyl transferase of forty species against \u003cem\u003eC. jejuni\u003c/em\u003e ADP-heptose-lipooligosaccharide heptosyl transferase exhibited that, out of forty species, only four species showed a significant identity percentage, including \u003cem\u003eC. coli\u003c/em\u003e, which showed 83% (e-value 0.0) identity; \u003cem\u003eC. taniopygae\u003c/em\u003e, which showed 78.5% ( e-value 2e-162) identity; \u003cem\u003eC. ornithocola\u003c/em\u003e, which represented 77.47% (e-value 4e-74) identity; and \u003cem\u003eC. lari\u003c/em\u003e, which showed 76.71% (e-value 4e-69) identity. However, thirty six species exhibited no significant similarity with \u003cem\u003eC. jejuni\u003c/em\u003e ADP-heptose-lipooligosaccharide heptosyl transferase. Apart from this, CMP-Neu5Ac synthase was found to be present in thirty species and CMP-N-acetylneuraminate-beta-galactosamide-alpha-2, 3-sialyltransferase was present in thirty two species.\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\u003eTypes of lipopolysaccharides and lipooligosaccharide synthesis enzymes in \u003cem\u003eCampylobacter\u003c/em\u003e species\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"10\"\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\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eS.No.\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eCampylobacter\u003c/em\u003e species\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eLipid A\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eOuter core\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eInner core\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eO-antigen\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eType of lipopolysaccharides (LPS)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eADP-heptose-lipooligosaccharide heptosyl transferase\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c9\"\u003e\u003cp\u003eCMP-Neu5Ac synthase\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c10\"\u003e\u003cp\u003eCMP-N-acetylneuraminate-beta-galactosamide-alpha-2,3-sialyltransferase\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. jejuni\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. coli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. fetus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. armoricus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003epresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. aviculae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. blaseri\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. canadensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePressent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. cuniculorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\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=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. curvus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. estrildidarum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. gracillis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. helveticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e13\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. hyointestinalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. mucosalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. Pinnipediorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. portucalensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. showae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. sputorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e19\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. subantracticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. troglodytis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. upsaliensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. ureolyticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. volucris\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e24\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. vulpis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e25\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. taeniopygae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e26\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. avium\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e27\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. concisus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. corcagiensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e29\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. geochelonsis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. hepaticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e31\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. hominis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e32\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. iguaniorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e33\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. insulanigrae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e34\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. laninae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. lari\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. novaezeelandiae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. ornithocola\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e38\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. pleoridis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e39\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. rectus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e40\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eCandidates C. infans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e41\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. analoticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. bilis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e43\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. cryaerophila\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. devanensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e45\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. magnus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. majalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. massiliensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e48\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. procelli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e49\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. suis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eDeep rough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eC. vicugnae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003eAbsent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eRough LPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c9\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c10\"\u003e\u003cp\u003ePresent\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section2\"\u003e\u003ch2\u003eGenome plasticity analysis\u003c/h2\u003e\u003cp\u003eHorizontal gene transfer (HGT) is a critical mechanism in the evolution of microbial genomes. In numerous bacterial pathogens, genomic islands and other mobile genetic elements facilitate movement of genetic repertoire and have been implicated in genome evolution via HGT (Juhas et al. 2015). Consequently, genomic islands were identified in twenty-nine species including, \u003cem\u003eC. armoricus, C. coli, C. avium, C. blaseri, C. canadensis, C. consisus, C. corcagiensis, C. cuniculorum, C. curvus, C. fetus, C. gracillis, C. helveticus, C. hominis, C. hyointestinalis, C. iguaniorum, C. laninae, C. lari, C. mucosalis, C. ornithocola, C. pinnipediorum, C. rectus, C. showae, C. sputorum, C. subantracticus, C. upsaliensis, Candidatus C. infans, C. procelli\u003c/em\u003e and \u003cem\u003eC. vicugnae\u003c/em\u003e. Certain genes associated with virulence were mapped to these genomic islands. These findings suggest that some virulence factors in these species may have been acquired through HGT (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eGenes associated with virulence and pathogenicity mapped to the genomic islands among \u003cem\u003eCampylobacter\u003c/em\u003e spp.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"2\"\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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGenes mapped to the genomic islands\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. fetus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType II toxin-antitoxin system, YafQ family toxin and type II toxin-antitoxin system RaE/ParE\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. coli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType VI secretion system protein, Type VI secretion system associated protein, type II CRISPR RNA guided endonuclease Cas9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. subantracticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType VI secretion system associated lipoprotein, Type VI secretion lysozyme-like protein\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. iguaniorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLPS biosynthesis protein, toxin-antitoxin system protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. gracilis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ehost nuclease inhibitor protein Gam, pathogenicity locus\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. rectus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDNA type IV secretion system protein ComB1a, type IV secretary system conjugative DNA transfer family protein, CRISPR associated protein, type VI secretion system\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. laninae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTetM/ TetW/ Tet O/ Tet S family tetracycline resistance ribosomal protection protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. procelli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cem\u003eCytolethal distending toxin CdtB\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. helveticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType IV secretion system and \u003cem\u003ecytolethal distending toxin subunit A, B\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. cuniculorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType IV secretion system protein virB9 was present.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. avium\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType II secretion system protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. canadensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType II secretion system protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hyointestinalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType IV secretary system conjugative\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. blaseri\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType IV secretion system DNA-binding domain-containing protein, virulence protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCandidates C. infans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eType II and type III secretion system protein, type I restriction endonuclease, type IV secretion system, virulence protein.\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eIt has been established that prophages play key role in HGT (Mancini et al. \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, Gomathinayagam et al. 2025). Among the 50 species of \u003cem\u003eCampylobacter\u003c/em\u003e, prophage regions were identified in 27 species, out of which 9 species harbored intact prophages (complete detail for supplementary table 2 (online resource) and Table \u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Most of the genes incorporated in the prophage region were annotated as hypothetical.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eGenomic islands, prophages, crisper arrays and spacers among the spp. of \u003cem\u003eCampyloabacter\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=\"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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eGenomic Islands\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eIntact prophages\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eCrisper arrays\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eSpacers\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. jejuni\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. coli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. fetus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e89\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\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e44\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. armoricus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e12\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. aviculae\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. blaseri\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. canadensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\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\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e42\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. cuniculorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e29\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. curvus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\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\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. estrildidarum\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. gracillis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e95\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. helveticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hyointestinalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e21\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. mucosalis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. Pinnipediorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. portucalensis\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. showae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. sputorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. subantracticus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. troglodytis\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. upsaliensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. ureolyticus\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. volucris\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. vulpis\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. taeniopygae\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. avium\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e2\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. concisus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. corcagiensis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. geochelonsis\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e78\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hepaticus\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. hominis\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e71\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. iguaniorum\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. insulanigrae\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. laninae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e18\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. lari\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e10\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. novaezeelandiae\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. ornithocola\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e6\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. pleoridis\u003c/em\u003e\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\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. rectus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e100\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCandidates C. infans\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. analoticus\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. bilis\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. cryaerophila\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. devanensis\u003c/em\u003e\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\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. magnus\u003c/em\u003e\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\u003e9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e10\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. majalis\u003c/em\u003e\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\u003e1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. massiliensis\u003c/em\u003e\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\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e46\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. procelli\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e12\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\u003e5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. suis\u003c/em\u003e\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\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. vicugnae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4\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\u003e2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e20\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThe CRISPR-Cas system is an adaptive immune system in bacteria and archaea that captures and stores sequences from invading elements (phages, plasmids) as spacers. Out of 50 species, twenty-six species were found to harbor CRISPR arrays (complete detail supplementary table 3 (online resource)) and Table \u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. A total of seven hundred spacers were identified in \u003cem\u003eCampylobacter\u003c/em\u003e spp., which denotes frequent infection by bacteriophages. Identification of spacers on the viral database revealed the prevalence of \u003cem\u003ePodoviridae\u003c/em\u003e and \u003cem\u003eMyoviridae\u003c/em\u003e as the major phages infecting \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section2\"\u003e\u003ch2\u003eResistome analysis\u003c/h2\u003e\u003cp\u003eAntibiotic resistance genes in \u003cem\u003eCampylobacter\u003c/em\u003e species pose a substantial public health challenge, as these bacteria are frequently implicated in foodborne illnesses. The presence of such genes complicates therapeutic interventions and may lead to increased morbidity among affected individuals. An analysis of antibiotic resistance genes across 50 \u003cem\u003eCampylobacter\u003c/em\u003e species revealed that 28 species possess genes conferring resistance to antibiotics (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). However, antibiotic resistance genes were not identified in 22 species including \u003cem\u003eC. fetus\u003c/em\u003e and \u003cem\u003eC. hepaticus\u003c/em\u003e which are pathogenic.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eAntibiotic resistance genes present in \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\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\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSpecies\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAntibiotic resistance drug class\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAMR gene Family\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eResistance mechanism\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. jejuni, C. coli, C. lari, C. volucris, Candidates C. infans, C. magnus\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePenicillin beta-lactam,\u003c/p\u003e\u003cp\u003emacrolide antibiotic, fluoroquinolone antibiotic, cephalosporin, fusidane antibiotic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eOXA beta-lactamase, OXA-61-like beta-lactamase,\u003c/p\u003e\u003cp\u003eresistance-nodulation-cell division (RND) antibiotic efflux pump\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAntibiotic inactivation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. portucalensis, C. armoricus, C. avium, C. blaseri, C. concisus, C. corcagiensis, C. geochelonsis, C. hyointrstinalis, C. ornithocola, C. pleoridis, C. rectus, C. showae, C. subantracticus, C. ureolyticus, C. analoticus, C. cryaerophila, C. devanensis, C. majalis, C. massiliensis, C. vicugnae.\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eFluoroquinolone antibiotic, tetracycline antibiotic,\u003c/p\u003e\u003cp\u003eglycopeptide antibiotic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eResistance-nodulation-cell division (RND) antibiotic efflux pump, glycopeptide resistance gene cluster, vanT\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAntibiotic efflux, antibiotic target alteration\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. magnus, C. taeniopygae, C. laninae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eTetracycline antibiotic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTetracycline-resistant ribosomal protection protein\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAntibiotic target protection\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eC. taeniopygae\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eAminoglycoside antibiotic\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eAPH(3')\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003eAntibiotic inactivation\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eComparative genomic analysis of prokaryotes has enhanced our understanding of the biology of various pathogenic microorganisms. The \u003cem\u003eCampylobacter\u003c/em\u003e genus includes a group of bacteria known for their significant role in causing infections in both humans and animals (Igwaran \u0026amp; Okoh, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Several \u003cem\u003eCampylobacter\u003c/em\u003e species have been known to cause serious infections in humans, underscoring their clinical significance. Over the last decade, the global incidence of campylobacteriosis has risen, indicating that it has become a widespread infectious disease (Bukari et al. \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). In \u003cem\u003eCampylobacter\u003c/em\u003e species, variations in genome size and the presence of plasmids are crucial factors that influence their adaptability, pathogenicity, and resistance to antimicrobials. Most \u003cem\u003eCampylobacter\u003c/em\u003e spp. have a relatively small genome, typically ranging between 1.4 to 2.9 Mb, indicating their streamlined metabolic capabilities and reliance on host environments. Despite its compact genome, \u003cem\u003eCampylobacter\u003c/em\u003e displays significant genetic diversity owing to frequent horizontal gene transfers and recombination. Plasmids were found to be harbored by nine spp. of \u003cem\u003eCampylobacter\u003c/em\u003e in this study, which greatly contribute to this diversity by carrying genes linked to virulence. For example, genes on plasmids of \u003cem\u003eC. fetus, C. pleoridis, C. helveticus\u003c/em\u003e and \u003cem\u003eC. lari\u003c/em\u003e encoding type IV secretion systems can enhance survival in challenging environments and promote gene exchange between species. Apart from that, the gene encoding cag pathogenicity island protein (cag12) was annotated on the plasmids of \u003cem\u003eC. pleoridis, C. helveticus\u003c/em\u003e and \u003cem\u003eC. lari\u003c/em\u003e. Therefore, genome size and plasmid content are one of the factors associated with the determination of the ecological fitness of \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003eThe phylogenomic relationships elucidated in this study offer a comprehensive understanding of the connections between pathogenic and non-pathogenic species within the genus, revealing that pathogenic species are related to some nonpathogenic or minor pathogenic species. Phylogenetic and phylogenomic analyses of \u003cem\u003eCampylobacter\u003c/em\u003e species have shown intriguing clustering patterns, where major pathogenic species, such as \u003cem\u003eC. jejuni\u003c/em\u003e and \u003cem\u003eC. coli\u003c/em\u003e, are closely grouped with minor or non-pathogenic species. This close association underscores the high degree of genetic relatedness within the genus and suggests that differences in pathogenicity of the spp. may stem from subtle variations in gene content, mobile genetic elements, and regulatory mechanisms, rather than broad genomic divergence. The clustering of pathogenic species with minor pathogenic members, such as \u003cem\u003eC. hepaticus\u003c/em\u003e, \u003cem\u003eC. upsaliensis\u003c/em\u003e or \u003cem\u003eC. hyointestinalis\u003c/em\u003e, indicates that the evolutionary paths of \u003cem\u003eCampylobacter\u003c/em\u003e are shaped by gene acquisition and loss events, horizontal gene transfer, and niche-specific adaptations.\u003c/p\u003e\u003cp\u003eComparative genome analysis based on metabolic pathways demonstrated a close relationship among \u003cem\u003eC. coli, C. bilis\u003c/em\u003e, and \u003cem\u003eC. jejuni\u003c/em\u003e, indicating that these species share a conserved metabolic framework that supports their survival and adaptation in diverse host environments. Clustering with \u003cem\u003eC. bilis\u003c/em\u003e, in which pathogenicity has not been reported, yet, highlights possibility that core metabolic pathways are conserved across pathogenic and non-pathogenic members of the genus. The close metabolic relatedness highlights a common evolutionary origin and functional similarity in nutrient utilization and energy metabolism, however, the absence of pathogenic traits in \u003cem\u003eC. bilis\u003c/em\u003e despite metabolic similarity to \u003cem\u003eC. jejuni\u003c/em\u003e and \u003cem\u003eC. coli\u003c/em\u003e underscores the importance of virulence repertoires in pathogenicity of the species rather than metabolic capacity alone.\u003c/p\u003e\u003cp\u003eEven though the species of \u003cem\u003eCampylobacter\u003c/em\u003e have been established as major or minor pathogens and been implicated in gastrointestinal infections, however, the information regarding the pathogenic potential of the rest of the species remains scarce. In this study, a genome-wide exploration of the factors associated with virulence was performed which provided key insights into virulence potential of non-pathogenic species. Based on the virulence repertoire, \u003cem\u003eC. vicugnae\u003c/em\u003e was clustered with C. \u003cem\u003ejejuni. C. vicugnae\u003c/em\u003e has been recently isolated from a patient suffering from gastroenteritis (Jehanne et al. \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), however, its pathogenicity has not been established yet. This indicates that \u003cem\u003eC. vicugnae\u003c/em\u003e harbors a subset of virulence associated determinants despite its limited association with clinical infections which may suggest either the presence of cryptic or unexplored virulence potential in \u003cem\u003eC. vicugnae\u003c/em\u003e, or an evolutionary retention of virulence genes that are not actively expressed under natural conditions. Overall, this observation highlights the need for further functional and epidemiological studies to assess whether \u003cem\u003eC. vicugnae\u003c/em\u003e could act as an opportunistic pathogen under certain conditions, or whether its virulence gene repertoire primarily represents evolutionary remnants without significant pathogenic outcomes. Some virulence factors associated with chemotaxis, adhesion and invasion play very important roles in the bacterial pathogenicity. High number of virulence factors (n\u0026thinsp;\u0026ge;\u0026thinsp;20) were observed in non pathogenic species (apart from major and minor pathogenic species) including \u003cem\u003eC. armoricus, C. avium, C. iguaniorum, C. laninae, C. ornithocola, C. showae, C. novaezeelandiae, C. volucris, C. subantracticus, C. taeniopygae, C. estrildidarum, C. aviculae, C. troglodytis, C. devanensis, C. vicugnae, C. vulpis, C. pleoridis, C. cuniculorum, C. procelli\u003c/em\u003e and \u003cem\u003eC. helveticus, C. insulanigare, C. mucosalis, C. rectus, C. sputorum\u003c/em\u003e, which have been associated with gastroenteritis and which have never been reported as pathogenic. The highest number of genes for virulence factors (n\u0026thinsp;=\u0026thinsp;29) were harbored by \u003cem\u003eC. jejuni\u003c/em\u003e, and \u003cem\u003eC. helveticus\u003c/em\u003e. \u003cem\u003eC. helveticus\u003c/em\u003e has been linked to gastroenteritis; however it has not been classified as a major or minor pathogen. The enrichment of adhesion, invasion, and toxin related genes in \u003cem\u003eC\u003c/em\u003e. \u003cem\u003ejejuni\u003c/em\u003e is consistent with its well established role as the leading cause of human campylobacteriosis, while the presence of similar virulence gene repertoire in \u003cem\u003eC. helveticus\u003c/em\u003e highlights its emerging significance as a potential pathogen. The higher virulence gene load may provide the species carrying them with a broader arsenal to overcome host defense mechanisms, establish persistent colonization, and cause disease. Such findings also indicate possible selective pressure or horizontal gene transfer events that have shaped the virulence architecture of these species.\u003c/p\u003e\u003cp\u003eThe process of chemotaxis involves the detection of external signals by bacteria, facilitating their movement towards favorable environments. This mechanism is employed by several pathogenic bacteria to invade their hosts. In \u003cem\u003eCampylobacter\u003c/em\u003e species, specifically in \u003cem\u003eC. jejuni\u003c/em\u003e, chemotaxis is mediated by glycoproteins and mucin, which serve as chemoattractants, guiding the bacteria to primary localization sites within the avian gut and the mucus-filled crypts of the avian ceca (Chang and Miller \u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The chemotaxis process utilizes two-component signal transduction pathways, involving six proteins: chemotaxis A, B, R, W, Y, and Z (Hermans et al. 2012), along with two methyl-accepting proteins. The CheW protein functions as a coupling protein, linking methyl-accepting proteins to CheA, which subsequently activates and transfers the phosphoryl group to either CheY or CheB. The binding of CheY proteins to the flagellar motor component FliM induces a change in rotation from counterclockwise to clockwise, thereby influencing bacterial swimming and motility. In addition to the six chemotaxis proteins, other proteins such as CetA, CetB, and LuxS are also implicated in the chemotaxis process in \u003cem\u003eCampylobacter\u003c/em\u003e species. Regarding adhesion as a virulence factor, Krause Craszczynska et al (2007) have previously reported that the \u003cem\u003eCadF\u003c/em\u003e gene is present in all \u003cem\u003eC. jejuni\u003c/em\u003e and \u003cem\u003eC. coli\u003c/em\u003e strains, is an outer membrane protein facilitating cell adhesion through its binding to the cell adhesion protein fibronectin. In present study, the \u003cem\u003eCadF\u003c/em\u003e gene was identified in 44 out of 50 \u003cem\u003eCampylobacter\u003c/em\u003e species. The colonization of \u003cem\u003eCampylobacter\u003c/em\u003e species in the host gut necessitates adherence to host gastrointestinal epithelial cells (Jin et al. \u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). Glycoproteins binding to fibronectin activate signaling GTPases Rac1 and Cdc42, which promote \u003cem\u003eCampylobacter\u003c/em\u003e cell internalization (Ziprin et al. \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). In terms of invasion, bacterial flagella play a crucial role during host invasion, serving as export apparatuses for the secretion of non-flagellar proteins (Guerry et al. 2007). The \u003cem\u003eFlac\u003c/em\u003e and \u003cem\u003eCia\u003c/em\u003e gene products utilize the flagellar secretion apparatus to be delivered into the host cell cytoplasm, which is essential for colonization and invasion into the host (Carrillo et al. \u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Among the 50 species of \u003cem\u003eCampylobacter\u003c/em\u003e, invasion genes \u003cem\u003eCiaB\u003c/em\u003e and \u003cem\u003eHtrA\u003c/em\u003e were identified 48 and 44 species respectively indicating invasion potential in these species.\u003c/p\u003e\u003cp\u003eThe cancer promoting potential of \u003cem\u003eC. jejuni\u003c/em\u003e in prolonged infections has been established and has been attributed to the cytolethal distending toxin produced by the species (Marroki et al. 2019; Zergui et al. \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The Cdt complex is composed of three subunits: A, B, and C. Thereby, all three subunits of \u003cem\u003eCdt\u003c/em\u003e genes were annotated in 22 species in this study. Apart from this, incomplete or non-functional \u003cem\u003eCdt\u003c/em\u003e genes missing either subunit A or subunit C genes were identified in \u003cem\u003eC. iguaniorum\u003c/em\u003e and \u003cem\u003eC. canadensis\u003c/em\u003e (no subunit C) and \u003cem\u003eC. avium, C. devanensis\u003c/em\u003e and \u003cem\u003eC. vicugnae\u003c/em\u003e (no subunit A). As per previous study, Cdt A and Cdt C attach to cholesterol-rich micro-domains on the cell membrane, facilitating the entry of Cdt B into cells via clathrin-coated pit endocytosis (Nesic et al. 2005). Once internalized Cdt B moves to the cytosol and may be transported retrogradely from the Golgi complex to the endoplasmic reticulum. Eventually, the Cdt B relocates to the nucleus, guided by a potential nuclear localization signal in its amino acid sequence. In the nucleus, Cdt B causes DNA double-strand breaks, leading to cell-cycle arrest (Lai et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Thus, Cdt can be considered as an important virulence marker as well as a key factor for promotion of tumor formation. The gene encoding Cdt was identified in all major (n\u0026thinsp;=\u0026thinsp;3) and 3 minor pathogenic species (out of 5) of the genus as well as 3 species (out of 6) which have been associated with gastroenteritis. However, the \u003cem\u003eCdt\u003c/em\u003e gene was also identified in 13 species in which pathogenicity has not been reported yet. Comparative analysis of the gene encoding \u003cem\u003eCdt\u003c/em\u003e in different \u003cem\u003eCampylobacter\u003c/em\u003e species with \u003cem\u003eCdt\u003c/em\u003e encoding gene of \u003cem\u003eC. jejuni\u003c/em\u003e, revealed that \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. vulpis\u003c/em\u003e (non-pathogenic) exhibit a high similarity to the \u003cem\u003eCdt\u003c/em\u003e gene of \u003cem\u003eC. jejuni\u003c/em\u003e. Furthermore, multiple sequence alignment of the amino acid sequences of Cdt B protein among different \u003cem\u003eCampylobacter\u003c/em\u003e species revealed conserved and variable regions that may contribute to their functional roles. The conservation of key amino acids such as histidine H152 and aspartate D185, D222 (except \u003cem\u003eC bilis\u003c/em\u003e) on catalytic sites across Cdt B protein sequences suggests evolutionary pressure to maintain structural and enzymatic function essential for Cdt activity. The presence of \u003cem\u003eCdt ABC\u003c/em\u003e gene in species not typically associated with disease suggests that possession of some virulence genes alone does not necessarily translate into virulence potential and may reflect an evolutionary remnant retained from a common ancestor or may have been acquired through horizontal gene transfer. Acquisition of other virulence factors in future might provide them with a complete arsenal for virulence. Phylogenetic analysis of the concatenated amino acid sequences of Cdt ABC revealed that \u003cem\u003eC. jejuni\u003c/em\u003e constitutes a distinct cluster, indicating unique sequence variations in comparison to other \u003cem\u003eCampylobacter\u003c/em\u003e species. This distinct separation highlights the evolutionary divergence of \u003cem\u003eCdt\u003c/em\u003e in \u003cem\u003eC. jejuni\u003c/em\u003e, which may contribute to its notable pathogenic potential. However, aside from the distinct clustering of \u003cem\u003eC. jejuni\u003c/em\u003e; \u003cem\u003eC. helveticus, C. upsaliensis\u003c/em\u003e, and \u003cem\u003eC. vulpis\u003c/em\u003e are closely related to \u003cem\u003eC. jejuni\u003c/em\u003e based on the amino acid sequences of \u003cem\u003eCdt\u003c/em\u003e, indicates a conserved evolutionary pattern and might have potential functional similarity of the toxin among these species.\u003c/p\u003e\u003cp\u003eThe secretion systems are essential for bacterial pathogenesis, adaptation, and interactions between host and pathogen. In \u003cem\u003eCampylobacter\u003c/em\u003e species, the T4SS and T6SS have become significant molecular mechanisms that enhance virulence, facilitate genetic exchange, and ensure survival in various environments. These systems, which are specialized protein complexes, enable bacteria to introduce effector proteins into eukaryotic (host cells) and prokaryotic (other bacteria) target cells. This injection can disrupt biological functions, enhance bacterial survival and reproduction within the host, and exacerbate disease (Wang et al. \u003cspan citationid=\"CR101\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Many bacteria, particularly gram-negative bacteria, possess the complex machinery known as T4SS. A set of VirB/VirD genes or their homologs, initially identified in \u003cem\u003eAgrobacterium tumefaciens\u003c/em\u003e, encode T4SS. T4SS typically includes an extracellular pilin composed of a minor VirB5 subunit and a major VirB2 subunit. Substrate secretion is driven by three ATPases, VirB4, VirB11, and VirD4, which may also assist in system assembly. The inner membrane channel comprises the biotopic membrane proteins VirB8 and VirB10, as well as the polytopic membrane protein VirB6, according to biochemical and functional studies. The composition of the pore at the outer membrane, which allows the substrate to enter the extracellular milieu, remains unknown. The short lipoprotein VirB7 may be a component of the complex VirB9. However, neither protein possesses a transmembrane region that has been identified or predicted. Additionally, the roles of VirB1 and VirB3 in this complex remain unclear (Fronzes et al. \u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). The T4SS genes were found on the plasmid in some species of the genus, including \u003cem\u003eC. fetus, C. lari, C. helveticus\u003c/em\u003e, and C. \u003cem\u003epleoridis\u003c/em\u003e, among which which \u003cem\u003eC. pleoridis\u003c/em\u003e is a non-pathogenic species. Their plasmid encodes many proteins of T4SS, indicating that the plasmid-associated secretion system may play a role beyond virulence, potentially contributing to ecological fitness rather than direct pathogenicity. Among the fifty species of \u003cem\u003eCampylobacter\u003c/em\u003e, T4SS proteins were identified in 29, indicating a likelihood of their involvement in horizontal gene transfer, plasmid-related functions, and adaptation. However, the uneven presence of the T4SS proteins among species indicates differing functionality and evolutionary diversification within the genus. Moreover, as reported by Gabbert et al (\u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), the T4SS was absent in \u003cem\u003eC. jejuni\u003c/em\u003e. The impact of the absence of T4SS is limited to its reduced capacity to perform functions typically associated with classical T4SS, such as horizontal gene transfer via conjugation or the direct injection of effectors into host cells, as seen in other pathogens. Instead, \u003cem\u003eC. jejuni\u003c/em\u003e has adapted to utilize other secretion systems, specifically the Type III secretion system (T3SS) and the T6SS, to facilitate its pathogenesis (Tikhomirova et al. \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). T4SS proteins were identified in a subset of species, suggesting that this system is not universally conserved throughout the genus. Apart from that, generally, 13\u0026ndash;15 core genes form the cluster encoding T6SS. This system mimics an inverted phage tail by puncturing target cells with a contraction-based mechanism. The \u003cem\u003eHcp\u003c/em\u003e gene, linked to the T6SS, creates a tube-like structure that allows certain substances and effector carriers to be secreted. The protein VgrG resembles a spike and penetrates the target membrane to deliver effectors. \u003cem\u003eClpV\u003c/em\u003e is an ATPase necessary for the recycling of sheath components such as \u003cem\u003etssC\u003c/em\u003e. The sheath components are \u003cem\u003etssB\u003c/em\u003e, which contracts during secretion, and \u003cem\u003etssC\u003c/em\u003e, which pairs with \u003cem\u003etssB\u003c/em\u003e. The base plate component \u003cem\u003etssE\u003c/em\u003e anchors the sheath. The baseplate core assembly genes are \u003cem\u003etssH, tssG\u003c/em\u003e, and \u003cem\u003etssK\u003c/em\u003e. The \u003cem\u003etssH\u003c/em\u003e gene is associated with the disassembly of the sheath for recycling (Pisarz et al. \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). A membrane complex protein gene includes \u003cem\u003etssJ, tssL\u003c/em\u003e, and \u003cem\u003etssM\u003c/em\u003e. Our analysis of T6SS across 50 \u003cem\u003eCampylobacter\u003c/em\u003e species reveals insights into the distribution patterns of these systems. Among the 50 species, 21 contained T6SS proteins, with 5 being major or minor pathogens, 2 linked to gastroenteritis, and 14 species for which pathogenicity has not yet been reported. Conversely, T6SS is increasingly recognized as a widespread and versatile secretion system, contributing to both inter-bacterial competition and environmental adaptation. Our study indicates that T6SS proteins are unevenly distributed among different \u003cem\u003eCampylobacter\u003c/em\u003e species.\u003c/p\u003e\u003cp\u003ePrevious research has established that the lipopolysaccharides constituting the outer membrane components of gram-negative bacteria, are well-documented virulence factors due to their impermeability to various toxins and enzymes, including proteases, lysozyme, detergents, and some hydrophobic antibacterial agents (Matsuura et al. 2013; Zhao et al. 2016). LPS is composed of three primary parts: lipid A, the core region, and O-antigen (O-polysaccharide). LPS is released in large amounts as a result of the lysis of bacterial cells, which can become an endotoxin, leading to pathological conditions, including sepsis. Thus, LPS is a well-established bacterial virulence factor responsible for pathogenesis (Sidor et al. 2023). Prior studies have shown that deep-rough LPS is more susceptible to antibiotics and reduce pathogenicity by obstructing the ADP heptose pathway (Sidor et al. 2023). An analysis of the outer membrane lipopolysaccharides of 50 \u003cem\u003eCampylobacter\u003c/em\u003e species revealed that 41 species exhibited and deep rough LPS and 9 species exhibited rough LPS which is more resistant to antibiotics. Furthermore, the sialylated Foligosaccharide structure found on \u003cem\u003eC. jejuni\u003c/em\u003e species is known to enhance their potential for pathogenicity. These structures possess epitopes that closely resemble ganglioside epitopes present in human peripheral nerves. \u003cem\u003eC. jejuni\u003c/em\u003e is classified into two categories based on whether the sialic acid component is linked to the bacterial carbohydrate outer surface through the action of sialyltransferases Cst-II or Cst-III. These bacteria exhibit a highly pathogenic phenotype and possess the capability to cause severe colitis (Louwen et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). The sialyated lipooligosaccharide synthesis enzymes analysis indicates that enzyme ADP-heptose lipooligosaccharide (LOS) heptosyltransferase, found in \u003cem\u003eCampylobacter\u003c/em\u003e species, plays a vital role in the synthesis and structural alteration of LOS, a crucial element of the outer membrane. This enzyme facilitates the addition of heptose units from ADP-heptose to the developing LOS core, thereby enhancing the structural variety and functional intricacy of LOS across various \u003cem\u003eCampylobacter\u003c/em\u003e species. These alterations are linked to immune system evasion, resistance to serum, and interactions between the host and pathogen. Notably, differences in heptosyltransferase genes between pathogenic and non-pathogenic \u003cem\u003eCampylobacter\u003c/em\u003e species may determine the degree of LOS truncation or extension, thereby influencing bacterial virulence potential. The presence of the enzyme in forty one species of the genus indicates evolutionary preservation of LOS biosynthesis pathways, while species-specific variations may signify adaptive strategies to unique ecological niches or host environments. Apart from that, the detection of CMP-Neu5Ac synthase in \u003cem\u003eCampylobacter\u003c/em\u003e species underscores a crucial element of their sialic acid metabolism. In present study CMP-Neu5Ac synthase was found to be present in thirty species. This enzyme facilitates the activation of N-acetylneuraminic acid (Neu5Ac) by transforming it into CMP-Neu5Ac, which acts as a donor substrate for sialyltransferases that modify surface structures like lipooligosaccharides (LOS). In \u003cem\u003eC. jejuni\u003c/em\u003e and some related species, CMP-Neu5Ac synthase is pivotal in promoting the sialylation of LOS, a mechanism that imitates host gangliosides, aiding in immune evasion, persistence, and occasionally leading to post-infectious neuropathies such as Guillain\u0026ndash;Barr\u0026eacute; syndrome (P\u0026eacute;rez et al. 2018). Notably, the presence and distribution of CMP-Neu5Ac synthase differ among \u003cem\u003eCampylobacter\u003c/em\u003e species, with pathogenic strains more commonly possessing this gene than non-pathogenic species. This variation implies that acquiring and maintaining CMP-Neu5Ac synthase may offer a selective advantage in host-associated environments by boosting virulence and adaptation. Consequently, the presence of CMP-Neu5Ac synthase not only highlights metabolic diversity within the genus but also signifies its evolutionary significance in shaping host-pathogen interactions. Furthermore, the enzyme CMP-N-acetylneuraminate β-galactosamide α-2,3-sialyltransferase, found in thirty two species of \u003cem\u003eCampylobacter\u003c/em\u003e, is a vital component of the sialylation process, connecting sialic acid metabolism to alterations in surface structures. This enzyme uses CMP-Neu5Ac as a donor to attach sialic acid to the terminal galactose units of lipooligosaccharides (LOS), resulting in sialylated LOS formation. These changes are crucial for host-pathogen interactions, as they allow bacteria to mimic human gangliosides, aiding in immune system evasion and resistance to serum. In pathogenic strains such as \u003cem\u003eC. jejuni\u003c/em\u003e, the function of this sialyltransferase is closely linked to increased virulence, survival within the host, and the potential to cause post-infectious autoimmune conditions such as Guillain\u0026ndash;Barr\u0026eacute; and Miller Fisher syndromes. However, this enzyme is not consistently present across the genus; many non-pathogenic or less virulent species do not possess it. This uneven distribution indicates that the presence and maintenance of CMP-N-acetylneuraminate β-galactosamide α-2, 3-sialyltransferase offers a selective benefit specifically for pathogenic \u003cem\u003eCampylobacter\u003c/em\u003e, highlighting its role in adaptive evolution and host colonization.\u003c/p\u003e\u003cp\u003eHorizontal gene transfer (HGT) is a pivotal evolutionary mechanism in prokaryotic genomes, facilitating the exchange of genetic material between organisms that are not in a parent-offspring relationship. This process is a well-documented adaptation strategy observed in bacteria and archaea. Although the implications of HGT extend beyond pathogenic organisms, it is frequently associated with microbial antibiotic resistance and pathogenicity (Soucy et al. \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). A substantial component of HGT is mediated by genomic islands (GEIs), which are distinct DNA segments that vary among closely related strains. Some GEIs are mobile, while others are either immobile or have lost their mobility. These islands can integrate into the host chromosome, excise, and transfer to a new host through transformation, conjugation, or transduction. GEIs are vital in the evolution of diverse bacterial species, as they facilitate the dissemination of variable genes, including those conferring antibiotic resistance and virulence (Juhas et al. \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Genomic islands were detected in 29 \u003cem\u003eCampylobacter\u003c/em\u003e species. The genomic islands in most species of this genus contain genes crucial for pathogenicity, such as pathogenic loci, type IV and type VI secretion systems, toxin-antitoxin systems, genes associated with antibiotic resistance, virulence genes, and cytolethal distending toxin subunits B and A. Toxin-antitoxin system in \u003cem\u003eCampylobacter\u003c/em\u003e is genetic elements that help to maintain plasmid stability and contributes to bacterial survival under stress. This suggests that HGT might be an important source for the evolution of pathogenic potential in this genus. Additionally, analysis of mobile genetic elements revealed that prophage regions were present in 27 species of \u003cem\u003eCampylobacter\u003c/em\u003e, with 9 species harboring intact prophages. CRISPR arrays were detected in 25 species, comprising a total of 700 spacers, suggesting their potential role in regulating HGT and shaping genomic plasticity within the genus. Furthermore, analysis of antibiotic resistance genes revealed that most pathogenic species exhibit these genes, while some nonpathogenic species show only 50% or less similarity with antibiotic resistance genes, indicating that pathogenic species have developed adaptive potential against drugs. The ability of microorganisms to acquire antibiotic resistance is well-established. The overuse and indiscriminate application of clinically prescribed antibiotics, coupled with the ongoing development and spread of mobile genetic resistance elements, have led to the emergence of multidrug-resistant (MDR) and even extremely drug-resistant (XDR) bacterial pathogens over the past two decades. The most commonly used antimicrobials for treating campylobacteriosis, when clinical intervention is necessary, are fluoroquinolones and macrolides; however, these are becoming less effective against \u003cem\u003eCampylobacter\u003c/em\u003e. Antibiotic resistance genes were found in 28 out of 50 species of the genus, indicating a broad but uneven distribution of resistance within the group. The species that possess these resistance genes demonstrate their ability to withstand antimicrobial pressure, raising concerns for treatment options.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003e\u003cem\u003eCampylobacter\u003c/em\u003e represents a clinically and epidemiologically important genus. Some species of the genus are key enteric pathogens. Comparative genomics of the \u003cem\u003eCampylobacter\u003c/em\u003e genus provides comprehensive insights into its evolutionary complexity, pathogenic potential and adaptive strategies. Phylogenetic and phylogenomic analysis showed that several non-pathogenic or minor pathogenic species of \u003cem\u003eCampylobacter\u003c/em\u003e clustered with major pathogenic species, indicating shared genomic features that could confer potential for pathogenicity under certain evolutionary or environmental conditions. Functional repertoire profiling underscored the metabolic flexibility of the genus, whereas virulence gene analysis, including detailed analysis of the \u003cem\u003eCdt\u003c/em\u003e gene, indicated that \u003cem\u003eC. helveticus\u003c/em\u003e possesses the potential to emerge as a pathogen. \u003cem\u003eC. vicugnae\u003c/em\u003e clustered with \u003cem\u003eC. jejuni\u003c/em\u003e suggesting shared virulence traits though its pathogenicity not yet well established, and the analysis of \u003cem\u003eCdt\u003c/em\u003e gene revealed, high similarity between \u003cem\u003eC. vulpis\u003c/em\u003e and \u003cem\u003eC. jejuni\u003c/em\u003e, highlighting conserved toxin-associated genes despite the lack of confirmed pathogenicity in \u003cem\u003eC. vulpis\u003c/em\u003e. The characterization of outer membrane components emphasizes their role in host interactions and immune invasion. Furthermore, evidence of genome plasticity, including HGT, indicates a dynamic evolutionary process driving diversity and pathogenicity. The evolution of pathogenic genes among \u003cem\u003eCampylobacter\u003c/em\u003e species may occur through horizontal gene transfer. A number of genomic islands have been identified in \u003cem\u003eCampylobacter\u003c/em\u003e species, and key virulence genes such as (\u003cem\u003eCdt\u003c/em\u003e) been mapped onto these islands. Viral signatures such as integrated prophages and CRISPR elements were abundant in the genomes of \u003cem\u003eCampylobacter\u003c/em\u003e spp., highlighting the high frequency of phage attack and therefore of HGT. Resistome analysis revealed the widespread presence of antibiotic resistance genes, underscoring the clinical significance of \u003cem\u003eCampylobacter\u003c/em\u003e as an emerging multidrug-resistant pathogen. Overall, these findings provide an integrative framework that not only enhances our understanding of \u003cem\u003eCampylobacter\u003c/em\u003e evolution and pathogenicity, but also providing valuable insights to strengthen future surveillance and therapeutic interventions. Furthermore, future investigations integrating transcriptomic, proteomic, and host-pathogen interaction analyses will be crucial for validating these genomic predictions and identifying novel determinants of pathogenicity and resistance. Future research should also include molecular docking of the Cdt protein to facilitate rational drug design against cytolethal distending toxin.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eCompeting interests\u003c/h2\u003e\n\u003cp\u003eThe authors have no relevant financial or non financial interests to disclose.\u003c/p\u003e\n\u003ch2\u003eFunding\u003c/h2\u003e\n\u003cp\u003eNA\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eThis study was conceived and designed by CT. TY performed all the analyses. The manuscript was written by TY and CT.\u003c/p\u003e\n\u003ch2\u003eData Availability\u003c/h2\u003e\n\u003cp\u003eThe sequence data used in this study is publicly available on NCBI and was not generated in this study. 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FEMS Microbiol Rev. 33(2):376\u0026ndash;393. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1111/j.1574-6976.2008.00136.x\u003c/span\u003e\u003cspan address=\"10.1111/j.1574-6976.2008.00136.x\" 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":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Campylobacter, gastrointestinal infection, cytolethal distending toxin, cancer, virulence, horizontal gene transfer","lastPublishedDoi":"10.21203/rs.3.rs-7771926/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7771926/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cem\u003eCampylobacter\u003c/em\u003e species are major contributors to foodborne and waterborne zoonotic gastroenteritis. Several species, including \u003cem\u003eC. jejuni, C. coli, C. fetus, C. concisus, C. lari, C. hyointestinalis, C. upsaliensis\u003c/em\u003e, and \u003cem\u003eC. hepaticus\u003c/em\u003e, are established pathogens, while the pathogenic potential of other members remains unclear. This study presents a comparative genomics analysis of the fifty reported species of \u003cem\u003eCampylobacter\u003c/em\u003e genus, encompassing phylogenomic relationships, functional repertoire profiling, virulence genes, diversity of Cytolethal distending toxin gene (\u003cem\u003eCdt\u003c/em\u003e), outer membrane components, genome plasticity, and resistome characterization. Phylogenetic analyses revealed that \u003cem\u003eC. hepaticus, C. taniopygae\u003c/em\u003e, and \u003cem\u003eC. iguaniorum\u003c/em\u003e, traditionally considered non-pathogenic or minor pathogens, cluster with major pathogenic species, suggesting shared evolutionary features. Functional repertoire profiling indicated metabolic flexibility that supports environmental adaptability, while virulence profiling highlighted both conserved and species-specific determinants. Variation in \u003cem\u003eCdt\u003c/em\u003e genes and outer membrane components emerged as key factors in pathogenicity. Notably, \u003cem\u003eC. helveticus\u003c/em\u003e shows potential to emerge as a significant pathogen, whereas \u003cem\u003eC. vicugnae\u003c/em\u003e and \u003cem\u003eC. vulpis\u003c/em\u003e display close evolutionary relationships with \u003cem\u003eC. jejuni\u003c/em\u003e. Genome plasticity analyses identified horizontal gene transfer via genomic islands, prophage insertions, and CRISPR arrays, underscoring the dynamic evolution of virulence traits. Resistome characterization revealed widespread antimicrobial resistance genes, raising concerns about multidrug resistance and clinical management. Overall, this study provides an integrative framework to understand the evolutionary dynamics, virulence potential, and antimicrobial resistance of \u003cem\u003eCampylobacter\u003c/em\u003e, offering valuable insights for surveillance and therapeutic strategies.\u003c/p\u003e","manuscriptTitle":"Comparative genomics of the Campylobacter genus: Insights into phylogenomics, virulence, genome plasticity and resistome profiling","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-10-25 05:34:36","doi":"10.21203/rs.3.rs-7771926/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c2b00c75-ddbd-4e7f-ab1f-60302716c201","owner":[],"postedDate":"October 25th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2025-12-12T10:54:20+00:00","versionOfRecord":[],"versionCreatedAt":"2025-10-25 05:34:36","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7771926","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7771926","identity":"rs-7771926","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}