Diversity and Parasitological Traits of Myxozoan Parasites Infecting Various Mullidae Species Along the Mediterranean Coasts of Türkiye

Diversity and Parasitological Traits of Myxozoan Parasites Infecting Various Mullidae Species Along the Mediterranean Coasts of Türkiye | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Diversity and Parasitological Traits of Myxozoan Parasites Infecting Various Mullidae Species Along the Mediterranean Coasts of Türkiye Erkan ÖZDEMİR, Cem Tolga GÜRKANLI This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9538756/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract Myxozoa are globally distributed microscopic parasites that primarily infect fish but also occur in amphibians and, more rarely, reptiles, birds, mammals, and annelid worms. The aim of this study was to determine the diversity and parasitological characteristics of Myxozoa infecting Mullus barbatus , Mullus surmuletus , and Upeneus moluccensis along the Mediterranean coasts of Türkiye. For this purpose, 330 M. barbatus , 253 M. surmuletus , and 240 U. moluccensis specimens were collected from three localities representing the western (Muğla), central (Antalya), and eastern (Mersin) regions of the Turkish Mediterranean coast and were subjected to parasitological investigations. Based on morphological and molecular analyses, three myxozoan species- Kudoa mediterraneus n. sp., Myxobolus mullus n. sp., and Ortholinea mullusi -were identified, two of which represent novel species. Kudoa mediterraneus n. sp., infecting the trunk muscles mainly of M. barbatus , was identified as a new species. This species represents the second Kudoa species reported from the family Mullidae worldwide and the first record of a Kudoa species infecting Mullidae in the Mediterranean Sea. The second novel species identified in this study, Myxobolus mullus n. sp., infects the gall bladder of M. barbatus and M. surmuletus . This species represents the second Myxobolus species, after Myxobolus parvus , reported to infect M. barbatus worldwide. Ortholinea mullusi , originally described from M. barbatus in the Black Sea, was recorded as the third myxozoan species in the present study. It was observed in the gonad, urinary bladder, and kidney of three M. barbatus specimens. Unexpectedly, no myxosporean infections were detected in U. moluccensis . Goatfish Kudoa Ortholinea Myxobolus Mediterranean Türkiye Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction The family Mullidae comprises 101 species classified within six genera [ 1 ]. In the Mediterranean Sea, six mullid species have been reported: Mullus barbatus Linnaeus 1758, M. surmuletus Linnaeus 1758, Upeneus moluccensis Bleeker 1855, U. pori Ben-Tuvia and Golani 1989, Parupeneus forsskali Fourmanoir & Guézé, 1976 and Pseudupeneus prayensis Cuvier 1829 [ 2 ]. Of these, only M. barbatus and M. surmuletus are native to the Mediterranean Sea, whereas the remaining species are invasive Lessepsian migrants originating from the Red Sea [ 3 ]. To date, five of the six species (excluding P. prayensis ) recorded from the Mediterranean Sea have also been reported along the Turkish coasts [ 2 ]. These species, particularly Mullus barbatus , M. surmuletus , and Upeneus moluccensis , are of considerable economic importance; therefore, numerous studies investigating their parasite fauna have been conducted both in Türkiye and worldwide. As a result, a rich parasitic fauna-particularly Digenea-has been reported from mullid species [ 4 – 8 ]. However, records of Myxozoa (Cnidaria), the target parasite group of this study, infecting Mullidae are relatively scarce [ 9 – 15 ]. Myxozoa (Cnidaria) are microscopic metazoan obligate endoparasites characterized by extremely reduced body size, structural simplicity, and complex two-host life cycle involving an invertebrate (polychaete and oligochaete) as the definitive host and a vertebrate (fish, amphibians, rarely reptiles, birds and mammals) as the intermediate host [ 16 – 19 ]. As can be expected from a lineage that diverged from its ancestor in the distant past, Myxozoa exhibit a remarkable biological diversity, with more than 2600 species described to date-corresponding to approximately 18% of all known Cnidaria species-classified within 64 genera and 17 families [ 17 , 19 , 20 ]. And from Türkiye a total of 31 myxozoan species have been reported, the majority of which (29 species) were recorded from fish hosts along the Black Sea coasts [ 15 , 21 ]. As mentioned earlier, although, Myxozoan species have been reported from a wide variaty of fish hosts worldwide, including Türkiye, there are only a few records of myxozoans infecting members of the Mullidae. In Turkiye, a total of three Myxozoan species- Myxobolus parvus , Ortholinea orientalis and Ortholinea mullusi -have been recorded parasitizing mullid species in the Blacks Sea; however, no records are currently available from the Marmara Sea, Aegean Sea, or Mediterranean Sea [ 22 – 24 ]. Additionally, severeal other records from different parts of the world are also available: Unicapsula galeata Naidjenova & Zaika, 1970 from Pseudopeneus pleurotaenia in the Indian Ocean; Ceratomyxa sp. from Mullus surmuletus in the Ría de Arousa in the northwest of Spain; Kudoa lutjanus Wang, Huang, Tsai, Cheng, Tsai, Chen, Chen, Chu, Liaw, Chang et Chen, 2005 from Upeneus tragula in China, Upeneus pflugfelderi Alama-Bermejo et al. 2009 from Pseudupeneus prayensis along the Atlantic coast of Africa and an unidentified myxozoan parasite from Mullus barbatus in Blanes-Spain [ 9 – 12 , 14 ]. However, none of these studies provide information on the diversity of Myxozoa infecting mullid species in the eastern Mediterrenean Sea. In this context, the aim of this study is to reveal the diversity of Myxozoa infecting three economically important mullid species- Mullus barbatus , M. surmuletus and Upeneus moluccensis -along the Mediterranean coasts of Türkiye; in this way, the study will provide the first data from eastern Mediterranean on this subject. Materials and Methods Fish Sampling and Parasitological Examination Between January and December 2024, a total of 330 Mullus barbatus L., 253 Mullus surmelatus and 240 Upeneus moluccensis specimens were collected from local fishermen in three locations-Muğla (37° 12′ 55″ N, 28° 21′ 49″ E), Antalya (36° 14’ 38’’ N, 29° 59’ 8’’ E) and Mersin (36° 48′ 44″ N, 34° 38′ 29″ E)-representing the Mediterranean coast of Türkiye (Figure S1 ). The trunk and various internal organs of fish specimens were examined parasitologically using a phase-contrast light microscope (Nikon Eclipse 80i, Nikon Corp., Tokyo, Japan). Detailed measurements and digital visualization of the observed myxospores were performed with a digital camera (Nikon Digital Sight DS-2Mv) mounted on the microscope. Measurements for all myxozoa species were based on 30 fresh spores and are presented as mean with minimum–maximum values given in parentheses (Table 1 ). Terminology and definitions for morphological identification were applied in accordance with Lom and Dyková [ 25 ]. In addition, prevalence of infection was determined according to Bush et al [ 26 ]. Table 1 Site of infection, hosts, geographical localities, and spore dimensions (µm) of Kudoa mediterraneus n. sp., identified in the present study, together with those of related myxozoan species. Species Spore body Polar capsule Site of infection Host species Locality References length width thickness length width Kudoa mediterraneus n. sp. 5.35 (6.79–4.12) 8.24 (9.75–7.19) 5.98 (6.96–4.82) 2.07 (2.99–1.38) 1.35 (1.94–1.01) Muscles Mullus barbatus , M. surmelatus Turkish coasts of Medditerranean (This study) Kudoa iidae 6.6 (7.0–6.0) Max: 11.6 (12.3–11) Min: 9.2 (9.9–8.6) Max: 8.1 (8.9–7.6) Min: 7.8 (7.0-8.5) Lp: 2.1 (2.3–1.8) Sp: 1.9 (2.2–1.6) Lp: 1.7 (1.8–1.6) Ip: 1.3 (1.6-1.0) Sp: 1.1 (1.3–0.8) Trunk muscle Pentanemus quinquarius Southeast Atlantic Ocean,off African coast [ 12 ] Kudoa trifolia 9.04 ± 0.64 (10.46–7.42) - 3.66 (3.96–3.32) Lp: 1.01 ± 0.03 (0.96–1.07) Sp: 0.84 ± 0.07 (0.96 − 0.76) Lp: 0.93 ± 0.05 (0.99 − 0.82) Sp: 0.79 ± 0.07 (0.9 − 0.7) Connective tissue Liza aurata Liza ramada Spain [ 41 ] Kudoa unicapsula 5.0 ± 0.24 (5.5–4.7) Max:8.0 ± 0.33 (8.4–7.2) Min: 7.7 ± 0.53 (8.3–6.6) Max: 6.1 ± 0.39 (6.9–5.3) Min:5.5 ± 0.43 (6.3–4.7) Lp: 3.1 ± 0.15 (3.4–2.8) Li: 1.4 ± 0.08 (1.5–1.3) Ls: 0.8 ± 0.05 (0.9 − 0.8) Lp: 2.7 ± 0.11 (2.9–2.5) Li: 0.8 ± 0.06 (0.7–0.9) Ls: 0.7 ± 0.05 (0.8 − 0.7) Intestinal mesentery, intestine and pyloric caeca Liza ramada Liza aurata Spain [ 39 ] Kudoa parvibulbosa 5.7 (6.4–4.6) 6.9 (7.2–6.3) 5.9 (6.7–5.7) 2.2 (2.4–1.8) 1.2 (1.4-1.0) Trunk muscles Megalaspis cordyla South China Sea, off Guangdong, [ 40 ] Molecular Analyses From each of the Kudoa sp. and Myxobolus sp., one specimen was selected for molecular analyses. In contrast, all three specimens of Ortholinea sp. were subjected to phylogenetic analysis, as they were obtained from different organs (Tables S1, S2, S3). An Invitrogen PureLink® Genomic DNA Mini Kit (USA) was used for total genomic DNA isolation from the infected host tissues. Extracted genomic DNA was stored at -20 ºC prior to use. The small subunit of nuclear ribosomal DNA (18S rDNA hereafter) was used as genetic marker for phylogenetic reconstructions. The PCR amplification of the 18S rDNA was performed using primer set MyxospecF [ 27 ] / 18r [ 28 ] with the following condition; an initial denaturation at 95°C for 4 minutes, followed by 40 cycles of denaturation at 94°C for 1 minute, annealing at 51°C (-0.1°C/Cyc) for 1 minute, and extension at 72°C for 2 minutes. The procedure was completed with a final extension at 72°C for 10 minutes. For samples that did not yield amplification with the previously mentioned primers, a nested PCR approach was employed. The initial amplification was performed using the primers ERIB1/ERIB10 [ 29 ], followed by a second round of amplification with MyxospecF/Myx4r [ 30 ]. Unlike the previously described PCR condition, the annealing temperature for the first amplification was set at 55°C. In the second round of PCR, the annealing temperature was also 55°C, and the extension step was carried out for 1 minute 45 seconds. For all amplifications, a 50 µl PCR reaction was prepared using, genomic DNA (50 ng), 1.5 mM MgCl 2 , 1.25 U Taq polymerase (New England BioLabs), 2.5 mM dNTP mix (Thermo Scientific), 5 µl of 10X PCR buffer, 0.5 pmol (final con.) of each primer and ddH 2 O. PCR amplification was carried out using a Techne (TC-Plus) thermal cycler, and the resulting PCR products were visualized on ethidium bromide–stained agarose gels using a imaging system (Vilber Lourmat, France). Nucleotide sequencings were performed commercially by Macrogen-Europe (Amsterdam, the Netherlands) with the same primers used for PCR amplifications. The software BioEdit v 7.2.5 [ 31 ] was employed to assemble the sequences from both strands. The data sets for phylogenetic analyses were conducted according to the results of the BLAST (Basic Local Alignment Search Tool, https://blast.ncbi.nlm.nih.gov/Blast.cgi ) searches and available literature (Tables S1, S2 and S3). The multiple nucleotide sequence alignments were performed using ClustalX 2.1 [ 32 ], and the resulting alignments were subsequently checked and manually edited using BioEdit v 7.2.5. The best-fitting evolutionary model(s) for each dataset were identified using the Akaike Information Criterion (AIC, [ 33 ]) and the Bayesian Information Criterion (BIC) tests, as implemented in the jModelTest v0.1 software package [ 34 , 35 ]. The Maximum Likelihood (ML) and Bayesian Inference (BI) were employed to infer phylogenetic relationships among genotypes and to construct phylogenetic trees. The ML analyses were performed using PhyML 3.0 [ 34 ], under the evolutionary models selected according to the AIC and BIC criteria. The robustness of the phylogenetic relationships inferred from the ML trees were assessed using bootstrap analysis [ 36 , 37 ], with 1,000 replicates. The Bayesian Inference (BI) analyses were performed using MrBayes v. 3.2 [ 38 ]. The analyses were conducted for ten million generations (ngen = 2,000,000) using two independent runs, each comprising four simultaneous Markov Chain Monte Carlo (MCMC) chains (one cold and three heated chains; nchains = 4). Trees and model parameters were sampled every 1,000 generations (samplefreq = 1000). The first 25% of samples were discarded as burn-in (burninfrac = 0.25). The substitution model was set to GTR with a proportion of invariable sites and gamma-distributed rate variation among sites (lset nst = 6, rates = invgamma). Nucleotide sequence similarities between genotypes were calculated using BioEdit v 7.2.5. The novel SSU rDNA genotypes obtained in this study were deposited in GenBank under accession numbers XXXXXXXX-XXXXXXXX. Results In this study, a total of 330 specimens of Mullus barbatus L., 1758, 253 specimens of Mullus surmuletus L., 1758, and 240 specimens of Upeneus moluccensis Bleeker, 1885, collected from three different locations (Muğla, Antalya, and Mersin) along the Mediterranean coast of Türkiye, were examined for myxosporean infections. As a result, three different myxosporean species belonging to three distinct genera were identified in the infected M. barbatus and M. surmuletus specimens. However, no myxosporean infections were detected in any of the Upeneus moluccensis specimens examined. The taxonomic summaries, morphological characteristics, and infection indices of the myxosporean species identified in this study are presented below; Kudoa mediterraneus n. sp. Taxonomic summary Phylum Cnidaria Hatschek, 1888 Class Myxozoa Grasse, 1970 Subclass Myxosporea Bütschli, 1881 Order Multivalvulida Shulman, 1959 Family Kudoidae Meglitsch, 1960 Genus Kudoa Meglitsch, 1947 Species name Kudoa mediterraneus n. sp. Type host Mullus barbatus L., 1758 (Red mullet) and Mullus surmelatus L., 1758 (Striped red mullet) Type locality Coasts of Muğla (37° 12′ 55″ N, 28° 21′ 49″ E), Antalya (36° 14’ 38’’ N, 29° 59’ 8’’ E) and Mersin (36° 48′ 44″ N, 34° 38′ 29″ E), Mediterranean, Türkiye. Site of infection Musculature. Type material One holotype (E-5) and one paratype (E-12), fixed in 70% ethanol, were deposited in the Parasitological Collection of the Fatsa Faculty of Marine Sciences, Ordu University, Ordu-Türkiye. Nucleotide Sequences The 18S rDNA nucleotide sequence obtained in this study was deposited in GenBank under accession number XXXXXXXX (specimen E-5). Etymology The specific epithet ‘ mediterraneus ’ refers to the geographical region from which the species was described. Description of myxospores No plasmodium was observed. In apical view, the myxospores were somewhat stellate in shape, composed of four equal shell valves, with indistinct suture lines. The posterior poles were slightly angular. Four equal-sized, ovoid to drop-shaped polar capsules were present; however, the coils of the polar flaments were not visible (Fig. 1 a, c). In lateral view, the myxospores were typically flattened to slightly convex at the posterior pole, with a rounded anterior pole. The polar capsules are clearly teardrop-shaped (Fig. 1 b). Myxospores have the following dimensions (n = 30): spore body length 5.35 (6.79–4.12) µm, spore body width 8.24 (9.75–7.19) µm, thickness 5.98 (6.96–4.82) µm, polar capsule length 2.07 (2.99–1.38) µm, polar capsule width 1.35 (1.94–1.01) µm (Table 1 ). Molecular phylogenetic analyses As a result of nucleotide sequencing, 1319 bp of the 18S rDNA was obtained from the Kudoa specimen (E-5) selected for phylogenetic analyses. BLAST analysis indicated that this genotype was closely related to Kudoa unicapsula (AM490334; [ 39 ]), K. parvibulbosa (LC626079; [ 40 ]), K. trifolia (AM183300; [ 41 ]), and K. iidae (LC493822; [ 12 ]). Subsequently, a dataset was constructed that included these species together with members of closely related (sister) sub-lineages within the genus Kudoa (Table S1 ). The phylogenetic analyses were conducted over 1221 aligned nucleotide positions with 164 polymorphic sites (184 substitutional mutations). The AIC and BIC tests suggested the GTR + I+G (I:0.727, G: 0.685) and TPM3uf + I+G (I:0.738, G: 0.709) evolutionary models, respectively. The ML trees constructed under both models revealed similar topologies; however, the tree generated using the former model was supported by higher bootstrap values. The phylogenetic relationships inferred from the BI analysis were largely consistent with those obtained using the ML approach, with only minor differences that did not influence the phylogenetic placement of the newly identified genotype in the present study. In this study, the ML tree generated under the GTR + I+G model is presented due to its higher bootstrap support. Bootstrap values from the ML (GTR + I+G) analysis and posterior probability values from the BI analysis are shown on the tree (Fig. 2 ). In both phylogenetic trees generated using the ML and BI algorithms, the genotype obtained in this study (E-5) formed a lineage with K. trifolia , K. unicapsula and K. iidae , showing nucleotide sequence similarities of 99.1%, 99.1% and 98.8%, respectively. In the ML tree, however, several additional species, including K. parvibulbosa , K. aburakarae and K. javaensis , also placed in this lineage; nevertheless, their inclusion was not supported by significant bootstrap values (Fig. 2 ). In contrast, in the BI tree these three species formed a distinct lineage together with K. quadricornis and K. paraquadricornis , which was recovered as sister to the lineage containing the newly obtained genotype. In both cases, the phylogenetic relationships of the newly obtained genotype and K. trifolia , K. unicapsularis and K. iidae were consistent and supported with sufficient bootstrap and posterior probability values. Remarks When the spores of Kudoa mediterraneus n. sp. were compared with those of the phylogenetically closest species ( K. trifolia , K. unicapsularis , K. iidae , and K. unicapsula ) and the geographically closest species ( K. nilüferi , K. anatolica , and Kudoa dicentrarchi ), all of these species were found to share unequal-sized valves and polar capsules. In contrast, Kudoa mediterraneus n. sp. possesses four equal-sized valves and polar capsules. In particular, K. unicapsularis , K. iidae , and K. unicapsula show pronounced variations in spore morphology that clearly distinguish them from Kudoa mediterraneus n. sp. Among these species, K. iidae exhibits a tail-like tapering extension of one of the shell valves, K. unicapsula possesses one polar capsule that is significantly larger than the others, and K. trifolia exhibits three small and one extremely large shell valves. Therefore, these species are morphologically very different from Kudoa mediterraneus n. sp and are unlikely to be confused with it. The geographically related species, on the other hand, all have genetic data (18S rDNA sequences) available in GenBank, which are clearly distinct from those of Kudoa mediterraneus n. sp. Prevalence, infection site(s), infection time and geographical distribution of Kudoa mediterraneus n. sp. Out of 330 Mullus barbatus specimens, 16 were infected with Kudoa mediterraneus n. sp., corresponding to a prevalence of 4.9%. Additionally, this species was detected in one of 253 specimens of M. surmelatus examined, representing a prevalence of 0.4%. However, no infection was detected in any of the 240 Upeneus moluccensis specimens examined. All K. mediterraneus specimens were observed in the muscles of the host fishes, and no clinical signs were detected in the infected tissues. Fifteen of the sixteen K. mediterraneus specimens were detected in summer, whereas only one was found in autumn. Geographically, twelve of the sixteen K. mediterraneus specimens were detected from Mersin (the eastern Mediterranean coasts of Turkiye), and four were detected in Antalya (the central Mediterranean coasts of Turkiye). However, no K. mediterraneus infection was observed in Muğla (the western Mediterranean coasts of Turkiye). Myxobolus mullus n. sp. Taxonomic summary Phylum Cnidaria Hatschek, 1888 Class Myxozoa Grasse, 1970 Subclass Myxosporea Bütschli, 1881 Order Bivalvulida Shul’man, 1959 Family Myxobolidae Thélohan, 1892 Genus Myxobolus Bütschli, 1882 Species name Myxobolus mullus n. sp. Type host Mullus barbatus L., 1758 (Red mullet) and Mullus surmelatus L., 1758 (Striped red mullet) Type locality Coasts of Muğla (37° 12′ 55″ N, 28° 21′ 49″ E), Antalya (36° 14’ 38’’ N, 29° 59’ 8’’ E) and Mersin (36° 48′ 44″ N, 34° 38′ 29″ E), Mediterranean, Türkiye. Site of infection Gall bladder Type material Holotype (E-2) and one paratype (E-17), fixed in 70% ethanol, were deposited in the Parasitological Collection of the Fatsa Faculty of Marine Sciences, Ordu University, Ordu-Türkiye. Nucleotide Sequences The 18S rDNA nucleotide sequence obtained in this study was deposited in GenBank under accession number XXXXXXXX (specimen E-2). Etymology The specific epithet ‘ mullus ’ refers to the host fish genus from which this parasite species was described. Description of vegetative stage and myxospores No plasmodia were observed. Fresh myxospores detected in the gall bladders were oval to oblong in valvular view and elliptical in sutural and apical views (Fig. 3 a-f). The measurements of the spore body were as follows (n = 30); 9.99 (11.11–9.05) µm in length, 7.23 (7.97–6.71) µm in width and 5.98 (6.53–5.46) µm in thickness. The two slightly unequally sized polar capsules were pyriform in shape and positioned parallel to each other. The larger polar capsule measured 3.87 (4.46–3.16) µm in length, 2.54 (2.91–2.13) µm in width, whereas the smaller polar capsule measured 3.53 (4.19-3.0) µm in length, 2.36 (2.84–2.02) µm in width (Table 2 ). Table 2 Site of infection, hosts, geographical localities, and spore dimensions (µm) of Myxobolus mullus n. sp. identified in the present study, together with those of related myxozoan species. Species Spore body Polar capsule Site of infection Host species Locality References length width thickness length width Myxobolus mullus n. sp. 9.99 (11.11–9.05) 7.23 (7.97–6.71) 5.98 (6.53–5.46) Lp: 3.87 (4.46–3.16) Ls: 3.53 (4.19-3.00) Lp: 2.54 (2.91–2.13) Ls: 2.36 (2.84–2.02) Gall Bladder Mullus Barbatus Turkish Coast of Mediterranean (This Study) Myxobolus iwagiensis 12.1 (12.8–11.3) 9.8 (10.5–9.1) 7.9 (8.2–7.5) Lp: 6.4 (6.9–5.9) Ls: 5.8 (6.5–5.3) Lp: 4.0 (4.4–3.4) Ls: 3.5 (4.0-3.1) Brain Oryzias latipes Japan [ 42 ] Myxobolus tai 10.2 (10.8–9.5) 7.6 ± 0.3 (8.1–7.1) 5.7 ± 0.2 (6.0-5.5) Lp: 4.4 (4.9–3.8) Ls: 4.0 ± 0.2 (4.5–3.6) Lp: 2.6 ± 0.1 (3.0-2.4) Ls: 2.6 ± 0.1 (2.9–2.3) Brain Pagrus major Japan [ 43 ] Myxobolus asymmetricus 10–11 6.5-7 5 - - Kidney connective tissue Symphodus tinca Italy [ 44 ] Myxobolus asymmetricus 8–10 5.8–6.5 - 4.4 - Kidney Labridae sp. Ukraine [ 45 ] Myxobolus asymmetricus 8.9–10.8 (10.1) 5.6–6.9 (6.5) - 4.4–5.3 (4.9) 4.4–5.3 (4.9) Kidney Parablennius tentacularis Turkey: Sinop (Black Sea) [ 23 ] Myxobolus asymmetricus 9.0–10.7 (7.1) 5.1–7.0 (4.3) - 4.5–5.3 (5.0) 2.9–3.4 (3.1) Kidney Parablennius sanguinolentus Turkey: Sinop (Black Sea) [ 23 ] Molecular phylogenetic analyses A 1585 bp fragment of the 18S rDNA gene was obtained from the Myxobolus sp. specimen (E-2) selected for phylogenetic analyses. The 18S rDNA genotypes of two recently described species, Myxobolus iwagiensis Kawano, Sakurai & Yanagida, 2025 (LC861743; [ 42 ]) and Myxobolus tai Kawano, Nitta & Yanagida, 2026 (LC884833, LC886104; [ 43 ]), were identified in BLAST analysis as the most similar sequences to the genotype newly obtained. Accordingly, a dataset comprising these two species and other closely related myxozoan species was constructed for further phylogenetic analyses (Table S2). However, the dataset was limited to 889 bp, including 436 variable sites and 659 mutations, due to the short length of some sequences and the high number of insertion and deletion mutations. The GTR + I+G (I:0.231, G: 0.592) and TIM2 + I + G (I:0.228, G: 0.578) evolutionary models were selected based on the AIC and BIC tests, respectively. The ML trees constructed under both models showed similar topologies, although higher bootstrap values were obtained under the first model. The BI analysis, conducted under GTR + I+G mode, produced a tree with a topology similar to that of the ML tree, showing only minor variation in the interlineage relationships within the first lineage. Due to its higher bootstrap support, the ML tree constructed with the GTR + I+G model is presented in Fig. 4 . Additionally, bootstrap values from the ML (GTR + I+G) analysis and posterior probability values from the BI analysis are indicated on the tree. In the phylogenetic trees generated using the ML and BI algorithms, three similar lineages were observed, one of which (the third lineage) comprised the newly obtained genotype in this study (E-2), together with Myxobolus tai and Myxobolus iwagiensis (Fig. 4 ). On both trees, the newly obtained genotype in this lineage appeared as sister taxa to M. tai genotypes, and this relationship was supported by 68% bootstrap value and 0.92 posterior probability. The nucleotide sequence similarity between E-2 and M. tai genotypes were 94.4%. The third species of the lineage, M. iwagiensis , placed as sister taxon to the previously mentioned species with 100% bootstrap support and 1.0 posterior probability. Additionally, the nucleotide sequence similarity between E-2 and M. iwagiensis genotype was 92.7%. Remarks Up to date, a total of nine Myxobolus species have been reported from the Turkish coasts, four of which possess unequal-sized polar capsules similar to those observed in the specimens obtained in the present study. However, three of these species have available genetic data (18S rDNA sequences) in GenBank that do not match the genotypes obtained in this study. One species, Myxobolus asymmetricus Parasi, 1912, lacks genetic data and therefore can only be compared based on morphological characteristics. When the spore measurements of Myxobolus mullus n. sp. are compared with those of M. asymmetricus and the two phylogenetically closest species, Myxobolus iwagiensis and Myxobolus tai , both the spore body and polar capsule measurements are significantly smaller than those of M. iwagiensis (Table 2 ) [ 23 , 44 , 45 ]. Furthermore, the two polar capsules of M. iwagiensis were markedly unequal in size, whereas those of M. mullus n. sp. were only slightly unequal. Although the spore body measurements of M. tai and M. mullus n. sp. are quite similar, the polar capsules of M. tai are considerably longer (Table 2 ). Finally, while the spore body lengths of M. asymmetricus and M. mullus n. sp. are quite similar, the spore body width is significantly greater in the M. mullus n. sp. (Table 2 ). Prevalence, infection site(s), infection time and geographical distribution of Myxobolus mullus n. sp. Myxobolus mullus n. sp. was detected in 23 of 330 Mullus barbatus specimens, corresponding to a prevalence of 7.0%. Additionally, the species was observed in 27 of 253 M. surmelatus specimens, corresponding to a prevalence of 10.7%. However, no infection with M. mullus n. sp. was detected in Upeneus moluccensis . All specimens of M. mullus n. sp. observed in both M. barbatus and M. surmelatus were recovered from the gall bladder. However, no clinical signs were observed in any the infected host fishes. Seasonally, majority of the M. mullus n. sp. infections ( n : 44) were detected during the spring and summer periods, whereas a few infections ( n : 6) were recorded in the winter and autumn periods. Geographically, of the 50 M. mullus n. sp. specimens, 35 were detected from Mersin (the eastern Mediterranean coasts of Turkiye), 10 were detected in Antalya (the central Mediterranean coasts of Turkiye), and 5 were detected in Muğla (the western Mediterranean coasts of Turkiye). Ortholinea mullusi Gürkanlı, Okkay, Çiftçi, Yurakhno and Özer 2018. Taxonomic summary Phylum Cnidaria Hatschek, 1888 Class Myxozoa Grassé, 1970 Subclass Myxosporea Bütschli, 1881 Order Bivalvulida Shul’man, 1959 Family Ortholineidae Lom & Noble, 1984 Genus Ortholinea Shul’man, 1962 Species name Ortholinea mullusi Gürkanlı, Okkay, Çiftçi, Yurakhno & Özer, 2018 Type host Mullus barbatus L., 1758 Type locality Coast of Sinop, Black Sea, Türkiye (42° 02’ 51’’ N, 35° 02’ 56’’ E). New locality Coasts of Muğla (37° 12′ 55″ N, 28° 21′ 49″ E) and Antalya (36° 14’ 38’’ N, 29° 59’ 8’’ E), Mediterranean, Türkiye. Site of infection Urinary bladder and kidney tubules. New site(s) of infection Gonads, Kidney and Urinary bladder Nucleotide Sequences The 18S rDNA nucleotide sequences obtained in this study were deposited in GenBank under accession numbers XXXXXXXX (specimen E-7), XXXXXXXX (specimen E-9) and XXXXXXXX (specimen E-13). Description of myxospores No plasmodium was observed. In frontal view, mature spores were spherical with rounded anterior and posterior poles (Fig. 5 a). However, in sutural and apical views, the spores were ellipsoidal (Fig. 5 b, c). Two polar capsules were equal in size and somewhat spherical. Notably, no polar flaments were observed. The dimensions of myxospores (n = 30 from each host) were as follows: spore body length 9.16 (9.52–8.92), width 8.33 (9.16–7.83), polar capsul length 3.16 (3.25–2.93) and polar capsul width 2.50 (2.62–2.41) (Table 3 ). Table 3 Site of infection, hosts, geographical localities, and spore dimensions (µm) of Ortholinea mullusi identified in the present study, together with those of related myxozoan species. Species Spore body Polar capsule Site of infection Host species Locality References length width thickness length width Ortholinea mullusi 9.16 (8.92–9.52) 8.33 (7.83–9.16) - 3.16 (2.93–3.25) 2.50 (2.41–2.62) Gonads, kidney, urinary bladder Mullus barbatus Muğla and Antalya coasts of the Mediterranean, Türkiye (This study) Ortholinea mullusi 9.3 (9.0 − 9.7) 8.7 (8.2 − 9.3) 7.7 (7.5 − 7.9) 3.1 (3.0-3.2) 2.5 (2.4–2.6) Urinary bladder, kidney Mullus barbatus Sinop coasts of the Black Sea, Türkiye [ 24 ] Molecular phylogenetic analyses Nucleotide sequencings yielded approximately 850–1700 bp of the 18S rDNA gene from the three Ortholinea specimens (E-7, E-9 and E-13), which were subsequently subjected to phylogenetic analyses. All three specimens exhibited identical 18S rDNA sequences. BLAST analyses revealed Ortholinea mullusi (MF539825; [ 24 ]) as the closest species to all newly obtained Ortholinea specimens (Table S3). For phylogenetic analyses, a dataset including O. mullusi and related Ortholinea species was constructed. The final alignment comprised 832 nucleotide positions, of which 281 were polymorphic (409 substitutional mutations). The AIC and BIC tests suggested GTR + I+G (I:0.39, G: 0.49) and TPM2uf + I+G (I:0.379, G: 0.46), evolutionary models, respectively. Similar phylogenetic relationships were obtained in the ML analyses conducted under both models. Likewise, the BI analysis produced a tree topology largely consistent with those inferred from the ML analysis. The ML tree generated under the GTR + I+G model is presented here due to its higher bootstrap support. Posterior probability values from the BI analysis are shown at the corresponding nodes together with ML bootstrap values (Fig. 6 ). In both ML and BI trees, the newly obtained Ortholinea genotype (E-7) clustered with O. mullusi , showing 99.3% nucleotide sequence similarity. This relationship was strongly supported by 97% bootstrap and a posterior probability of 1.0. Ortholinea sp. specimen RT-1 (MK937851; [ 46 ]) appeared as a sister to this lineage, with 54% bootstrap support and a posterior probability of 0.81. Remarks Comparison of the three Ortholinea specimens obtained in this study with O. mullusi , their closest phylogenetic relative, revealed a high degree of morphological similarity of the myxospores in frontal (spherical), sutural (ellipsoidal), and apical (spherical) views (Fig. 5 a-c). Likewise, the spor body length and width as well as the polar capsul length and width, of the newly obtaned Ortholinea specimens were fully consistent with those of O. mullusi (Table 3 ). The only difference between them was the shape of the polar capsules: in the original description of O. mullusi , the polar capsules were pyriform and equal in size, whereas in the newly obtained specimens, they were also equal in size but nearly ovoidal in shape. Prevalence, infection site(s), infection time and geographical distribution of Ortholinea mullusi Only three out of 330 examined M. barbatus specimens were found to be infected with O. mullusi , corresponding to a prevalence of 0.9%. However, no infection was detected in M. surmelatus and U. moluccensis specimens. All three O. mullusi infections were observed in different host organs including, the gonads, urinary bladder and kidney. Additionally, no clinical signs were detected in the infected organs. Two of the three O. mullusi infections were recorded in host specimens collected from Antalya (the central Mediterranean coasts of Turkiye) in summer, while one was detected in autumn from Muğla (the the western Mediterranean coasts of Turkiye). Discussion In the Mediterranean Sea, the family Mullidae, which includes economically important species, comprises six species, five of which have also been reported from the Mediterranean coasts of Türkiye [ 2 ]. Despite their economic importance, there are only a few records concerning the myxozoan diversity infecting members of the Mullidae family, and none of them relate to the western Mediterranean [ 9 , 11 , 14 , 22 – 24 , 47 ]. So, this study provided new results on the myxozoan diversity infecting Mullus barbatus , Mullus surmuletus , and Upeneus moluccensis collected from three locations, Muğla, Antalya, and Mersin along the Mediterranean coasts of Türkiye. Initial microscopic examinations revealed specimens belonging to three different myxozoan genera, Kudoa Meglitsch, 1947, Myxobolus Bütschli, 1882, and Ortholinea Shulman, 1962, in various organs of M. barbatus and M. surmuletus . However, no myxosporean infections were detected in Upeneus moluccensis specimens. The Kudoa infections were exclusively detected in the trunk muscles of M. barbatus ( n : 16) and M. surmuletus ( n : 1) specimens. The complete similarity in myxospor morphology, the concordance of morphometric data, and the shared source tissue (trunk muscles) indicate that all specimens belonged to the same Kudoa species (Fig. 1 a-c; Table 1 ). In the phylogenetic trees constructed using the ML and BI algorithms, the Kudoa sp. genotype obtained in this study (E-5) formed a distinct lineage together with Kudoa trifoli , Kudoa unicapsula , and Kudoa iidae , supported by high bootstrap values and posterior probabilities (Fig. 2 ). Within the genus Kudoa , interspecific nucleotide sequence similarity based on 18S rDNA is generally high compared with other myxozoan genera. For example, the similarity is 99.9% between Kudoa quadricornis and Kudoa paraquadricornis , 99.8% between Kudoa yasunagai and Kudoa chaetodoni , and 99.3% between Kudoa grammatorcyni and Kudoa scomberomori [ 48 ]. From this perspective, the nucleotide sequence similarities between the new genotype identified in the present study and the above-mentioned species (ranging from 99.1% to 98.8%) are not sufficiently high to assign the new genotype to any of these previously described species. This assumption was further supported by the myxospore morphology and morphometric data (Table 1 ). All phylogenetically related species- K. trifolia , K. unicapsula , and K. iidae -possess unequal shell valves and polar capsules [ 12 , 39 , 41 ]. In contrast, the specimens obtained in the present study exhibited morphologically homogeneous spores with equally sized shell valves and polar capsules, clearly distinguishing them from the aforementioned species. Based on the morphological, morphometric, and molecular phylogenetic data, we propose that the newly obtained Kudoa specimens represent a new species, designated as Kudoa mediterraneus n. sp. The genus Kudoa currently includes 134 binomial species. Of these, three species have so far been reported from the Turkish seas: K. dicentrarchi Sitja-Bobadilla & Alvarez-Pellitera, 1992 (ex Dicentrarchus labrax ), K. anatolica Özer, Okkay, Gürkanlı, Yılmaz, Yurakhno, 2018 (ex Atherina hepsetus ) and K. nilüferi Özer, Okkay, Gürkanlı, Yılmaz, Yurakhno, 2018 (ex Neogobius melanostomus ) [ 48 , 49 ]. Accordingly, K. mediterraneus n. sp. represents the forth species of Kudoa recorded from the Turkish coasts. So far, species of the genus Kudoa have been reported to infect a wide variety of fish species across different families [ 50 ]. However, to the best of our knowledge, there has been only one record of a Kudoa species, K. lutjanus Wang, Huang, Tsai, Cheng, Tsai, Chen, Chen, Chu, Liaw, Chang & Chen, 2005, infecting a member of the Mullidae, specifically Upeneus tragula in China [ 14 ]. Therefore, Kudoa mediterraneus n. sp. is the second Kudoa species described from a host in the family Mullidae worldwide. Additionally, it also represents the first Kudoa species known to infect mullid hosts in the Mediterranean Sea. The genus Kudoa , which forms the histozoic lineage of Myxozoa, has been mostly found in the skeletal muscle of fish hosts. Additionally, infections have also been reported in tissues of various other organs, including the gills, brain, heart, kidney, spleen, ovary, gall bladder, urinary bladder, oesophagus, intestine, mesentery, and smooth muscle [ 51 , 52 ]. Consistent with this, all Kudoa mediterraneus specimens in this study were determined in the trunk muscles. Although some Kudoa species, such as K. thyrsites , K. musculoliquefaciens , and K. clupeidae , have been linked to post-harvest softening of fish flesh, no such pathological effects have been observed for K. mediterraneus [ 53 – 55 ]. Species of the genus Kudoa have been reported to reach peak infection rates during summer and autumn [ 56 – 58 ]. In full agreement with this pattern, sixteen of the seventeen Kudoa mediterraneus specimens were found in summer, with only one specimen observed in autumn. The prevalence of K. mediterraneus was 4.9% (16/330) in M. barbatus and 0.4% (1/253) in M. surmuletus , while no infection was detected in the examined U. moluccensis specimens examined. When compared to prevalences reported for other Kudoa species from the Mediterranean and adjacent seas (the Aegean Sea, Sea of Marmara, and Black Sea)—including K. unicapsula from Liza ramada (40%) and Liza aurata (15%), K. camarguensis from Pomatoschistus minutus (2.02%) and Pomatoschistus microps (12.58%), K. nilüferi from Neogobius melanostomus (12.8%), and K. anatolica from Atherina hepsetus (32.1%)—the infection levels observed in the present study seem comparatively low [ 39 , 48 , 59 ]. Moreover, these values are much lower than the prevalence reported for K. lutjanus , the only Kudoa species previously documented from a mullid host, in U. tragula (20%) [ 14 ]. However, considering the prevalence of K. mediterraneus n. sp. in M. barbatus separately by location, it was 0.8%, 1.7%, and 13.3% in Muğla (western Mediterranean coast of Türkiye), Antalya (central Mediterranean coast of Türkiye), and Mersin (eastern Mediterranean coast of Türkiye), respectively. Additionally, only one K. mediterraneus specimen was found in M. surmuletus during the summer period in Antalya (one of 120 specimens examined, corresponding to a prevalence of 0.8%). These patterns suggest that M. barbatus might be the main host for this parasite, which could be more prevalent in the eastern Mediterranean and might still be spreading toward the western Mediterranean. The second and most abundant myxozoan specimens identified in this study belonged to the genus Myxobolus Bütschli, 1882, accounting for 50 infections among a total of 823 examined specimens of M. barbatus , M. surmuletus , and U. moluccensis . All observed Myxobolus specimens showed similar myxospore morphology and morphometric characteristics (Fig. 3 a-f; Table 2 ). Additionally, all specimens were found exclusively in the same organ, the gall bladder. All these data indicate that the specimens belong to the same Myxobolus species. In both ML and BI trees based on 18S rDNA nucleotide sequences, the newly obtained genotype (E-2) formed a lineage with Myxobolus tai [ 43 ] and M. iwagiens [ 42 ] that supported by significant bootstrap values and posterior probabilities (Fig. 4 ). Considering the 18S rDNA nucleotide sequence identities between some closely related Myxobolus species in our dataset, these values range from 96.9% to 98.2% (Table S4), indicating that sequence identities below the minimum threshold of 98.2% may correspond to distinct species. This threshold value for species delineation may be even higher when all Myxobolus species are considered. In the light of this data, the nucleotide sequence identities were insufficient to assign the specimens obtained in this study to either of M. tai and M. iwagiensis , as the sequence similarities between the new genotype (E-2) and M. tai and M. iwagiensis were 94.4% and 92.7%, respectively, which are well below the 98.2% threshold. The morphology and morphometric characteristics of the myxospores also supported the molecular findings. Although the newly obtained Myxobolus specimens and the two phylogenetically closest species, M. tai and M. iwagiensis , share a similar oblong spore shape, M. iwagiensis differs from the new specimens by having larger spore body dimensions, while M. tai has larger polar capsules compared to those of the new specimens [ 42 , 43 ]. Myxobolus asymmetricus (lacking molecular data), a species previously reported from Türkiye, shows significantly narrower polar capsules than the new specimens (Table 2 ) [ 23 , 44 , 45 ]. Finally, M. parvus , reported from M. barbatus in the Black Sea, differs from the new specimens by having equal-sized polar capsules [ 23 ]. Based on the combined morphological, morphometric, and molecular phylogenetic data, the examined Myxobolus specimens are described here as a new species, Myxobolus mullus n. sp. The genus Myxobolus Bütschli, 1882 is the most speciouse group within the phylum Myxozoa, comprising nearly 1000 valid species [ 60 , 61 ]. Of these, nine species- M. asymmetricus Lalitha-Kumari, 1969 (ex. Parablennius sanguinolentus and Parablennius tentacularis ), M. episquamalis Egusa, Maeno & Sorimachi (ex. Mugil cephalus ), M. exiguus (ex. Mugil cephalus ), M. ichkeulensis (ex. Mugil cephalus ), M. muelleri Bütschli, 1882 (ex. Mugil cephalus and Diplodus annularis ), M. parvum Yurakhno, 1991 (ex. Parablennius tentacularis ), M. parvus Shulman, 1962 (ex. Liza saliens , Chelon saliens and Mullus barbatus ), M. rotundus Nemeczek, 1911 (ex. Symphodus cinereus ) and M. spinacurvatura Maeno, Sorimachi, Ogawa & Egusa, 1990 (ex. Mugil cephalus )- have been reported from the Turkish coasts [ 13 , 15 , 23 , 62 – 66 ]. Moreover, six of these species have been reported from the Black Sea coasts of Türkiye, while two were recorded from the Mediterranean Sea and one from the Sea of Marmara. Therefore, M. mullus represents the tenth Myxobolus species reported from the Turkish coasts and the third Myxobolus species recorded from the Mediterranean coast of Türkiye. Moreover, as mentioned earlier, only a limited number of myxozoan species have been reported as parasites of the Mullidae family, one of which is M. parvus , described from the Black Sea [ 23 ]. Accordingly, M. mullus represents the second Myxobolus species reported from M. barbatus worldwide. Based on the infection characteristics of M. mullus n. sp., this species reached its peak infection rate (44 out of 50 cases) during the spring and summer seasons, when sea temperatures range from approximately 18°C to 28°C [ 67 ]. This pattern aligns with previously reported infection periods of Myxobolus species [ 68 – 70 ]. All fifty M. mullus specimens in this study were found exclusively in the gall bladder. However, the organ -or tissue- specificity of this species remains uncertain. Several Myxobolus species reported from the gall bladder— M. bankimi Sarkar, 1999; M. chilkensis Kalavati, Venkateswara Rao & Vaidahei, 1992; M. myleus Azevedo, Clemente, Casal, Matos, Alves, Al-Quraishy & Matos, 2012; M. hepatobiliaris Rocha, Casal, Alves, Antunes, Rodrigues & Azevedo, 2019; and M. vesicularis Rocha, Casal, Alves, Antunes, Rodrigues & Azevedo, 2019—have not been reported from other organs or tissues so far. Conversely, some species, such as M. cuneus Adriano, Arana & Cordeiro, 2006 and M. parvus , have been documented from various organs -including the urinary bladder, gills, spleen, fins, head surface, lower jaw, liver, heart, kidney tubules, and within several cysts- in addition to the gall bladder [ 60 , 71 ]. Therefore, further research is necessary to clarify this issue. Overall, the prevalence of M. mullus n. sp. in M. barbatus and M. surmuletus was 7% and 10.7%, respectively. These rates appear relatively low compared with previous reports of various Myxobolus species recorded in Türkiye, where prevalence values ranged from 9% to 53% (general around 20–25% for most species) [ 13 , 15 , 23 , 62 – 66 ]. However, when analyzed by sampling location, the prevalence of M. mullus n. sp. in Muğla (the western Mediterranean coast of Türkiye) was 12.5% in M. barbatus and 27.4% in M. surmuletus . In contrast, prevalence was only 3.3% and 5% in Antalya (central Mediterranean coast of Türkiye) for M. barbatus and M. surmuletus , respectively, and 4.4% and 1.7% in Mersin (eastern Mediterranean coast of Türkiye) (Table S5). In conclusion, 70% (35 out of 50) of all M. mullus n. sp. infections identified in this study were recorded from the western Mediterranean region. If the sea surface temperature factor is disregarded, since temperatures were similar across the sampling locations during the same months as mentioned in Şişman et al. [ 67 ], this pattern suggests that M. mullus n. sp. may still be spreading from the western Mediterranean to the eastern Mediterranean. Microscopic examination of three M. barbatus specimens (out of 330 examined) revealed a third myxozoan parasite belonging to the genus Ortholinea Shulman, 1962. However, this parasite was not observed in either M. surmuletus or U. moluccensis specimens. All three Ortholinea specimens (E-7, E-9, and E-13) exhibited an identical 18S rDNA genotype, clustering with Ortholinea mullusi (MF539825; Gurkanlı et al., 2018) in both ML and BI phylogenetic trees (Fig. 6 ). The 18S rDNA sequence similarity between O. mullusi and the newly obtained genotype was 99.3%. Rangel et al. [ 72 ] reported that interspecific variation in 18S rDNA within the genus Ortholinea ranges from 1.65% to 29.1%, suggesting that the isolates in this lineage are likely conspecific specimens of O. mullusi . Additionally, the close similarity in myxospore morphology and the agreement of morphometric data further support this conclusion (Table 3 ). The only difference observed between the original description of O. mullusi and the newly obtained specimens was the shape of the polar capsules, which were pyriform in the original description and slightly ovoidal in the new specimens. We believe this difference may be due to adaptation to different geographical locations, which show significant variation in certain physical parameters, such as sea temperature and salinity (renging from 15‰ to 20‰ in the Black Sea and approximatesly 39‰ in the Mediterranean) [ 73 , 74 ]. Therefore, based on morphological, morphometric, and molecular phylogenetic evidence, we assign the newly obtained Ortholinea specimens in this study to O. mullusi Gürkanlı, Okkay, Çiftçi, Yurakhno & Özer, 2018. The genus Ortholinea Shulman, 1962 currently includes 27 valid species, most of which infect marine fish hosts [ 72 ]. Only a few exceptions, such as O. fluviatilis , O. africanus , O. lauquen , and O. sphaerocapsularae , have been reported from non-marine environments [ 75 – 77 ]. Although the genus shows relatively low species diversity compared to other myxozoan genera, its members have a broad global distribution [ 78 ]. In Türkiye, five Ortholinea species have been reported to date: O. divergens (ex Parablennius sanguinolentus ), O. gobiusi (ex Neogobius melanostomus ), O. orientalis (ex Alosa tanaica and Mullus barbatus ), O. mullusi (ex Mullus barbatus ), and O. hamsiensis (ex Engraulis encrasicolus ) [ 22 – 24 , 78 ]. All of these species have been recorded from the Black Sea coasts of Türkiye. However, to date, no Ortholinea species have been reported from the Sea of Marmara or along the Turkish coasts of the Aegean and Mediterranean Seas. Ortholinea mullusi , the species identified in this study, was originally described from M. barbatus along the Black Sea coasts of Türkiye and has not been reported elsewhere [ 24 ]. Therefore, this study represents the second global record for this species. Given its occurrence in the Black Sea and the Mediterranean Sea, it is plausible that O. mullusi also infects M. barbatus populations in the Aegean and Marmara Seas. However, further research is needed to confirm this. Species of the genus Ortholinea are mainly coelozoic and have predominantly been reported from organs of the urinary system [ 72 ]. Consistent with these data, O. mullusi was originally described from the urinary bladder and kidney [ 24 ]. Similarly, in the present study, O. mullusi specimens E-9 and E-13 were observed in the kidney and urinary bladder, respectively. Additionally, specimen E-7 was detected in the gonad and the attached oviduct. Considering the coelozoic nature of the genus Ortholinea , it is more likely that this specimen originated from the oviduct rather than the gonadal tissue. Likewise, another species of the genus, O. undulans , has been reported from both the urinary system and the oviduct [ 79 ]. Gürkanlı et al. [ 24 ] reported an infection prevalence of O. mullusi in M. barbatus along the Black Sea coast of 24.5% (49 infected hosts out of 200 examined fish). In contrast, in this study, this parasite was detected in only 3 of the 330 examined M. barbatus individuals (0.9%). Moreover, as mentioned earlier, it was not observed in any specimens of M. surmuletus or U. moluccensis . Additionally, O. mullusi was detected in M. barbatus specimens collected from Muğla (the western Mediterranean coast of Turkiye) and Antalya (the central Mediterranean coast of Turkiye), but no infection was recorded in samples from Mersin (the eastern Mediterranean coast of Turkiye). The significant difference in infection rates between the Black Sea and Mediterranean populations, along with the limited distribution of the parasite in the Mediterranean Sea, suggests that O. mullusi may have only recently been introduced into the Mediterranean Sea. The species’ adaptation and expansion processes could still be ongoing. In this study, three mullid hosts were examined for myxozoan infections; however, no myxozoan parasites were observed in U. moluccensis . This host is a Lessepsian species, with the earliest records along the eastern Mediterranean coast dating back to the 1947 [ 80 ]. Although U. moluccensis has a wide native distribution, extending from the Red Sea, East Africa, Madagascar and Réunion east to the Caroline Islands and New Guinea, north to southern Japan, and southward to Western Australia and Queensland (Australia), there are no documented records of myxozoan infections for this species [ 81 ]. However, one myxozoan parasite, Ceratomyxa sultani , has been reported from another Upeneus species, U. margarethae , in the Arabian Gulf, suggesting that U. moluccensis may also harbor myxosporean parasites in its native range [ 82 ]. Upon migration to the eastern Mediterranean, this fish may have lost its myxosporean parasites, potentially due to the absence of the appropriate primary invertebrate host. Future studies incorporating various ecological factors are needed to further elucidate this phenomenon. The main outcomes of this study are summarised as follows: a) a new myxosporean species, named Kudoa mediterraneus n. sp., was identified from the trunk muscles of M. barbatus and M. surmuletus ; b) this species is the second Kudoa species known to infect the Mullidae family worldwide and the first Kudoa species recorded from Mullidae in the Mediterranean Sea; c) a second new myxosporean species, named Myxobolus mullus n. sp., was identified in the gall bladder of M. barbatus and M. surmuletus ; d) this species is the second Myxobolus species, after M. parvus , reported to infect M. barbatus globally; e) this study provides the first record of O. mullusi from the Mediterranean Sea; f) unexpectedly, no myxosporean infections were found in U. moluccensis . Declarations Competing interests The authors declare that they have no conflict of interest. Ethical standards All applicable international, national, and institutional guidelines for the care and use of animals were followed. Financial support This study received no grant from any funding agency. Author Contribution E. Özdemir provided support for the fieldwork of the study and initial laboratory studies, including dissections and parasitological examinations. For molecular work, C.T. Gürkanlı and E. Özdemir obtained DNA sequence data from parasites and conducted the phylogenetic analyses. C.T. Gürkanlı and E. Özdemir prepared the firs draft of the manuscript. C.T. Gürkanlı oversaw all subsequent revisions of the manuscript and wrote the final text. All authors reviewed the manuscript and provided edits on the final version. Acknowledgement The authors gratefully acknowledge Prof. Dr. Ahmet Özer for his valuable contributions to the finalization of the manuscript. Data availability statement Sequence data is available on the NCBI GenBank database. All other necessary data are included in the article. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9538756","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":637003510,"identity":"f15bbbdb-bc14-4d2b-9204-3b51387da6b2","order_by":0,"name":"Erkan ÖZDEMİR","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABCklEQVRIiWNgGAWjYHACAzBiYGAD4gogZmdgYGwgXssZBgYeZrgWZjxaGKBaGNuI0MLPf3jjoxsF9xj4pY8lfi6cZ5O4n5n54MMZDHZyug38xz5g0SLZcKzYOMegmEGyL+2w9MxtaYk9zGzJhhsYko3NDjAzz8DmqoM9ZtI5BgkMBmfYG6R5tx0GauExk3zAcCBxG1ALVo8c5jH/DdJif4a9+TfvHGK0HOMxYwbbwsN2TJq3AaplAx4tkj1sxSCH8UicYUuz5jmWZtxzGOiXGQZAvxxmNsYVYp9z/iTI8fewGd/mqbGRbW9vPviwp8JOzux442OsoQwFPOgOZsATk6NgFIyCUTAKCAEA3+NVr/ioGzoAAAAASUVORK5CYII=","orcid":"","institution":"Ordu University","correspondingAuthor":true,"prefix":"","firstName":"Erkan","middleName":"","lastName":"ÖZDEMİR","suffix":""},{"id":637003511,"identity":"ce283d19-1c64-4c14-b1dc-ca2b6179136d","order_by":1,"name":"Cem Tolga GÜRKANLI","email":"","orcid":"","institution":"Ordu University","correspondingAuthor":false,"prefix":"","firstName":"Cem","middleName":"Tolga","lastName":"GÜRKANLI","suffix":""}],"badges":[],"createdAt":"2026-04-27 08:38:49","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9538756/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9538756/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":109200491,"identity":"22429f0a-3c2f-4719-a446-7db221c2eff5","added_by":"auto","created_at":"2026-05-13 13:45:37","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":651117,"visible":true,"origin":"","legend":"\u003cp\u003eMature spore of \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. observed in the gall bladder in the present study \u003cstrong\u003ea)\u003c/strong\u003e apical view; \u003cstrong\u003eb)\u003c/strong\u003e lateral view; \u003cstrong\u003ec)\u003c/strong\u003e schematic drawing showing the details of a spore in apical, scale bar=10 µm.\u003c/p\u003e","description":"","filename":"OnlineFig.1.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/5a763c3b21cfc66c9ec0a71f.png"},{"id":109200497,"identity":"2941de72-3a6f-48db-9ca0-8c2c9988d2cb","added_by":"auto","created_at":"2026-05-13 13:45:43","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":526703,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum-likelihood (ML) phylogram showing the relationships of \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. genotypes (E-5) generated in the present study with related \u003cem\u003eKudoa\u003c/em\u003e species retrieved from the NCBI database (GenBank accession numbers in parentheses; see Table S1 for details). The tree was inferred under the GTR+I+G model and rooted using \u003cem\u003eKudoa thunni\u003c/em\u003e and \u003cem\u003eK. scomberi\u003c/em\u003e. Bootstrap values (≥50) from ML analysis and posterior probabilities (≥0.5) from BI analysis are indicated at the corresponding nodes.\u003c/p\u003e","description":"","filename":"OnlineFig.2.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/258a8f1292f0b9628090b4e1.png"},{"id":109200500,"identity":"6f5aa12c-d9d5-4b2f-9c22-9ec4d06bb850","added_by":"auto","created_at":"2026-05-13 13:45:44","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1681118,"visible":true,"origin":"","legend":"\u003cp\u003eMature spore of \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp. observed in the gall bladder in the present study \u003cstrong\u003ea)\u003c/strong\u003e valvular view; \u003cstrong\u003eb)\u003c/strong\u003e sutural view; \u003cstrong\u003ec)\u003c/strong\u003e apical view; \u003cstrong\u003ed, e, f)\u003c/strong\u003e schematic drawing showing the details of the spores in the valvular, sutural and apical views, respectively. Scale bar=10 µm\u003c/p\u003e","description":"","filename":"OnlineFig.3.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/4333688528b9ab9f08b2c87f.png"},{"id":109200501,"identity":"fe7377ae-a059-4a06-8d28-d0d26ca7c9c0","added_by":"auto","created_at":"2026-05-13 13:45:44","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":669468,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum-likelihood (ML) phylogram showing the relationships of \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp. genotypes (E-2) generated in the present study with related \u003cem\u003eKudoa\u003c/em\u003e species retrieved from the NCBI database (GenBank accession numbers in parentheses; see Table S2 for details). The tree was inferred under the GTR+I+G model and rooted using \u003cem\u003eHoferellus cyprini\u003c/em\u003eand \u003cem\u003eH. carassii\u003c/em\u003e. Bootstrap values (≥50) from ML analysis and posterior probabilities (≥0.5) from BI analysis are indicated at the corresponding nodes.\u003c/p\u003e","description":"","filename":"OnlineFig.4.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/f32ff6963adc751101f16623.png"},{"id":109200567,"identity":"159c7780-e3a8-4520-a23a-0cedc0846b00","added_by":"auto","created_at":"2026-05-13 13:46:18","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":3911427,"visible":true,"origin":"","legend":"\u003cp\u003eMature spore of \u003cem\u003eOrtholinea mullusi\u003c/em\u003e observed in the urinary bladder in the present study \u003cstrong\u003ea)\u003c/strong\u003e apical view; \u003cstrong\u003eb)\u003c/strong\u003e sutural view; \u003cstrong\u003ec)\u003c/strong\u003e apical view\u003c/p\u003e","description":"","filename":"OnlineFig.5.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/3a311f0e4af17c8acdfb7522.png"},{"id":109200498,"identity":"44cafef8-d025-4363-9852-829cb18a181f","added_by":"auto","created_at":"2026-05-13 13:45:43","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":422928,"visible":true,"origin":"","legend":"\u003cp\u003eMaximum-Likelihood (ML) tree illustrating the phylogenetic relationships between \u003cem\u003eOrtholinea\u003c/em\u003egenotypes obtained in this study (E-7, E-9 and E-13) and related Myxozoa species retrieved from NCBI database (GenBank accession numbers are given in parenthesis, see Table S3 for details). The tree was constructed under the GTR+I+G substitution model and rooted with \u003cem\u003eSphaerospora festivus\u003c/em\u003e. Bootstrap values (≥50) from ML analysis and posterior probabilities (≥0.5) from BI analysis are indicated at the corresponding nodes.\u003c/p\u003e","description":"","filename":"OnlineFig.6.png","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/a516a9e1263ea99c9eff737d.png"},{"id":109200710,"identity":"bcc8a222-2e80-480e-b8ef-598b9791eb1d","added_by":"auto","created_at":"2026-05-13 13:46:42","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":11522280,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/09963c9e-b685-40a8-afb1-e762ed063b31.pdf"},{"id":109200496,"identity":"823024dc-ddd4-4155-8b3e-efa764b061e6","added_by":"auto","created_at":"2026-05-13 13:45:42","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":506034,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial.docx","url":"https://assets-eu.researchsquare.com/files/rs-9538756/v1/db34e146f1491528ae8d27ef.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Diversity and Parasitological Traits of Myxozoan Parasites Infecting Various Mullidae Species Along the Mediterranean Coasts of Türkiye","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe family Mullidae comprises 101 species classified within six genera [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. In the Mediterranean Sea, six mullid species have been reported: \u003cem\u003eMullus barbatus\u003c/em\u003e Linnaeus 1758, \u003cem\u003eM. surmuletus\u003c/em\u003e Linnaeus 1758, \u003cem\u003eUpeneus moluccensis\u003c/em\u003e Bleeker 1855, \u003cem\u003eU. pori\u003c/em\u003e Ben-Tuvia and Golani 1989, \u003cem\u003eParupeneus forsskali\u003c/em\u003e Fourmanoir \u0026amp; Gu\u0026eacute;z\u0026eacute;, 1976 and \u003cem\u003ePseudupeneus prayensis\u003c/em\u003e Cuvier 1829 [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. Of these, only \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e are native to the Mediterranean Sea, whereas the remaining species are invasive Lessepsian migrants originating from the Red Sea [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. To date, five of the six species (excluding \u003cem\u003eP. prayensis\u003c/em\u003e) recorded from the Mediterranean Sea have also been reported along the Turkish coasts [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. These species, particularly \u003cem\u003eMullus barbatus\u003c/em\u003e, \u003cem\u003eM. surmuletus\u003c/em\u003e, and \u003cem\u003eUpeneus moluccensis\u003c/em\u003e, are of considerable economic importance; therefore, numerous studies investigating their parasite fauna have been conducted both in T\u0026uuml;rkiye and worldwide. As a result, a rich parasitic fauna-particularly Digenea-has been reported from mullid species [\u003cspan additionalcitationids=\"CR5 CR6 CR7\" citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. However, records of Myxozoa (Cnidaria), the target parasite group of this study, infecting Mullidae are relatively scarce [\u003cspan additionalcitationids=\"CR10 CR11 CR12 CR13 CR14\" citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eMyxozoa (Cnidaria) are microscopic metazoan obligate endoparasites characterized by extremely reduced body size, structural simplicity, and complex two-host life cycle involving an invertebrate (polychaete and oligochaete) as the definitive host and a vertebrate (fish, amphibians, rarely reptiles, birds and mammals) as the intermediate host [\u003cspan additionalcitationids=\"CR17 CR18\" citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. As can be expected from a lineage that diverged from its ancestor in the distant past, Myxozoa exhibit a remarkable biological diversity, with more than 2600 species described to date-corresponding to approximately 18% of all known Cnidaria species-classified within 64 genera and 17 families [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. And from T\u0026uuml;rkiye a total of 31 myxozoan species have been reported, the majority of which (29 species) were recorded from fish hosts along the Black Sea coasts [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. As mentioned earlier, although, Myxozoan species have been reported from a wide variaty of fish hosts worldwide, including T\u0026uuml;rkiye, there are only a few records of myxozoans infecting members of the Mullidae. In Turkiye, a total of three Myxozoan species-\u003cem\u003eMyxobolus parvus\u003c/em\u003e, \u003cem\u003eOrtholinea orientalis\u003c/em\u003e and \u003cem\u003eOrtholinea mullusi\u003c/em\u003e-have been recorded parasitizing mullid species in the Blacks Sea; however, no records are currently available from the Marmara Sea, Aegean Sea, or Mediterranean Sea [\u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. Additionally, severeal other records from different parts of the world are also available: \u003cem\u003eUnicapsula galeata\u003c/em\u003e Naidjenova \u0026amp; Zaika, 1970 from \u003cem\u003ePseudopeneus pleurotaenia\u003c/em\u003e in the Indian Ocean; \u003cem\u003eCeratomyxa\u003c/em\u003e sp. from \u003cem\u003eMullus surmuletus\u003c/em\u003e in the R\u0026iacute;a de Arousa in the northwest of Spain; \u003cem\u003eKudoa lutjanus\u003c/em\u003e Wang, Huang, Tsai, Cheng, Tsai, Chen, Chen, Chu, Liaw, Chang et Chen, 2005 from \u003cem\u003eUpeneus tragula\u003c/em\u003e in China, \u003cem\u003eUpeneus pflugfelderi\u003c/em\u003e Alama-Bermejo et al. 2009 from \u003cem\u003ePseudupeneus prayensis\u003c/em\u003e along the Atlantic coast of Africa and an unidentified myxozoan parasite from \u003cem\u003eMullus barbatus\u003c/em\u003e in Blanes-Spain [\u003cspan additionalcitationids=\"CR10 CR11\" citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. However, none of these studies provide information on the diversity of Myxozoa infecting mullid species in the eastern Mediterrenean Sea. In this context, the aim of this study is to reveal the diversity of Myxozoa infecting three economically important mullid species-\u003cem\u003eMullus barbatus\u003c/em\u003e, \u003cem\u003eM. surmuletus\u003c/em\u003e and \u003cem\u003eUpeneus moluccensis\u003c/em\u003e-along the Mediterranean coasts of T\u0026uuml;rkiye; in this way, the study will provide the first data from eastern Mediterranean on this subject.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eFish Sampling and Parasitological Examination\u003c/h2\u003e \u003cp\u003eBetween January and December 2024, a total of 330 \u003cem\u003eMullus barbatus\u003c/em\u003e L., 253 \u003cem\u003eMullus surmelatus\u003c/em\u003e and 240 \u003cem\u003eUpeneus moluccensis\u003c/em\u003e specimens were collected from local fishermen in three locations-Muğla (37\u0026deg; 12\u0026prime; 55\u0026Prime; N, 28\u0026deg; 21\u0026prime; 49\u0026Prime; E), Antalya (36\u0026deg; 14\u0026rsquo; 38\u0026rsquo;\u0026rsquo; N, 29\u0026deg; 59\u0026rsquo; 8\u0026rsquo;\u0026rsquo; E) and Mersin (36\u0026deg; 48\u0026prime; 44\u0026Prime; N, 34\u0026deg; 38\u0026prime; 29\u0026Prime; E)-representing the Mediterranean coast of T\u0026uuml;rkiye (Figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). The trunk and various internal organs of fish specimens were examined parasitologically using a phase-contrast light microscope (Nikon Eclipse 80i, Nikon Corp., Tokyo, Japan). Detailed measurements and digital visualization of the observed myxospores were performed with a digital camera (Nikon Digital Sight DS-2Mv) mounted on the microscope. Measurements for all myxozoa species were based on 30 fresh spores and are presented as mean with minimum\u0026ndash;maximum values given in parentheses (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Terminology and definitions for morphological identification were applied in accordance with Lom and Dykov\u0026aacute; [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. In addition, prevalence of infection was determined according to Bush et al [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e].\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSite of infection, hosts, geographical localities, and spore dimensions (\u0026micro;m) of \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp., identified in the present study, together with those of related myxozoan 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=\"char\" char=\".\" 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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c4\" namest=\"c2\"\u003e \u003cp\u003eSpore body\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003ePolar capsule\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSite of infection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eHost species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eReferences\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ethickness\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.35 (6.79\u0026ndash;4.12)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.24 (9.75\u0026ndash;7.19)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.98 (6.96\u0026ndash;4.82)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.07 (2.99\u0026ndash;1.38)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.35 (1.94\u0026ndash;1.01)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eMuscles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMullus barbatus\u003c/em\u003e, \u003cem\u003eM. surmelatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTurkish coasts of Medditerranean\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e(This study)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKudoa iidae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e6.6 (7.0\u0026ndash;6.0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMax: 11.6 (12.3\u0026ndash;11)\u003c/p\u003e \u003cp\u003eMin: 9.2 (9.9\u0026ndash;8.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMax: 8.1 (8.9\u0026ndash;7.6)\u003c/p\u003e \u003cp\u003eMin: 7.8 (7.0-8.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 2.1 (2.3\u0026ndash;1.8)\u003c/p\u003e \u003cp\u003eSp: 1.9 (2.2\u0026ndash;1.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 1.7 (1.8\u0026ndash;1.6)\u003c/p\u003e \u003cp\u003eIp: 1.3 (1.6-1.0)\u003c/p\u003e \u003cp\u003eSp: 1.1 (1.3\u0026ndash;0.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTrunk muscle\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003ePentanemus quinquarius\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSoutheast Atlantic Ocean,off African coast\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKudoa trifolia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e9.04\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64 (10.46\u0026ndash;7.42)\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\u003e3.66 (3.96\u0026ndash;3.32)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 1.01\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03 (0.96\u0026ndash;1.07)\u003c/p\u003e \u003cp\u003eSp: 0.84\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 (0.96\u0026thinsp;\u0026minus;\u0026thinsp;0.76)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 0.93\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 (0.99\u0026thinsp;\u0026minus;\u0026thinsp;0.82)\u003c/p\u003e \u003cp\u003eSp: 0.79\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07 (0.9\u0026thinsp;\u0026minus;\u0026thinsp;0.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eConnective tissue\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eLiza aurata\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003cem\u003eLiza ramada\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSpain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKudoa unicapsula\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.24 (5.5\u0026ndash;4.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMax:8.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33 (8.4\u0026ndash;7.2)\u003c/p\u003e \u003cp\u003eMin: 7.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53 (8.3\u0026ndash;6.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMax: 6.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39 (6.9\u0026ndash;5.3)\u003c/p\u003e \u003cp\u003eMin:5.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43 (6.3\u0026ndash;4.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 3.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15 (3.4\u0026ndash;2.8)\u003c/p\u003e \u003cp\u003eLi: 1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08 (1.5\u0026ndash;1.3)\u003c/p\u003e \u003cp\u003eLs: 0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 (0.9\u0026thinsp;\u0026minus;\u0026thinsp;0.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 2.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11 (2.9\u0026ndash;2.5)\u003c/p\u003e \u003cp\u003eLi: 0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06 (0.7\u0026ndash;0.9)\u003c/p\u003e \u003cp\u003eLs: 0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05 (0.8\u0026thinsp;\u0026minus;\u0026thinsp;0.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eIntestinal mesentery, intestine and pyloric caeca\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eLiza ramada\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003cem\u003eLiza aurata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSpain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eKudoa parvibulbosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e5.7 (6.4\u0026ndash;4.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.9 (7.2\u0026ndash;6.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.9 (6.7\u0026ndash;5.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2.2 (2.4\u0026ndash;1.8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e1.2 (1.4-1.0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eTrunk muscles\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMegalaspis\u003c/em\u003e\u003c/p\u003e \u003cp\u003e\u003cem\u003ecordyla\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSouth China Sea,\u003c/p\u003e \u003cp\u003eoff Guangdong,\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]\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\n\u003ch3\u003eMolecular Analyses\u003c/h3\u003e\n\u003cp\u003eFrom each of the \u003cem\u003eKudoa\u003c/em\u003e sp. and \u003cem\u003eMyxobolus\u003c/em\u003e sp., one specimen was selected for molecular analyses. In contrast, all three specimens of \u003cem\u003eOrtholinea\u003c/em\u003e sp. were subjected to phylogenetic analysis, as they were obtained from different organs (Tables S1, S2, S3). An Invitrogen PureLink\u0026reg; Genomic DNA Mini Kit (USA) was used for total genomic DNA isolation from the infected host tissues. Extracted genomic DNA was stored at -20 \u0026ordm;C prior to use. The small subunit of nuclear ribosomal DNA (18S rDNA hereafter) was used as genetic marker for phylogenetic reconstructions. The PCR amplification of the 18S rDNA was performed using primer set MyxospecF [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e] / 18r [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e] with the following condition; an initial denaturation at 95\u0026deg;C for 4 minutes, followed by 40 cycles of denaturation at 94\u0026deg;C for 1 minute, annealing at 51\u0026deg;C (-0.1\u0026deg;C/Cyc) for 1 minute, and extension at 72\u0026deg;C for 2 minutes. The procedure was completed with a final extension at 72\u0026deg;C for 10 minutes. For samples that did not yield amplification with the previously mentioned primers, a nested PCR approach was employed. The initial amplification was performed using the primers ERIB1/ERIB10 [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e], followed by a second round of amplification with MyxospecF/Myx4r [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. Unlike the previously described PCR condition, the annealing temperature for the first amplification was set at 55\u0026deg;C. In the second round of PCR, the annealing temperature was also 55\u0026deg;C, and the extension step was carried out for 1 minute 45 seconds. For all amplifications, a 50 \u0026micro;l PCR reaction was prepared using, genomic DNA (50 ng), 1.5 mM MgCl\u003csub\u003e2\u003c/sub\u003e, 1.25 U Taq polymerase (New England BioLabs), 2.5 mM dNTP mix (Thermo Scientific), 5 \u0026micro;l of 10X PCR buffer, 0.5 pmol (final con.) of each primer and ddH\u003csub\u003e2\u003c/sub\u003eO. PCR amplification was carried out using a Techne (TC-Plus) thermal cycler, and the resulting PCR products were visualized on ethidium bromide\u0026ndash;stained agarose gels using a imaging system (Vilber Lourmat, France). Nucleotide sequencings were performed commercially by Macrogen-Europe (Amsterdam, the Netherlands) with the same primers used for PCR amplifications. The software BioEdit v 7.2.5 [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e] was employed to assemble the sequences from both strands. The data sets for phylogenetic analyses were conducted according to the results of the BLAST (Basic Local Alignment Search Tool, \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://blast.ncbi.nlm.nih.gov/Blast.cgi\u003c/span\u003e\u003cspan address=\"https://blast.ncbi.nlm.nih.gov/Blast.cgi\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) searches and available literature (Tables S1, S2 and S3). The multiple nucleotide sequence alignments were performed using ClustalX 2.1 [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e], and the resulting alignments were subsequently checked and manually edited using BioEdit v 7.2.5. The best-fitting evolutionary model(s) for each dataset were identified using the Akaike Information Criterion (AIC, [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]) and the Bayesian Information Criterion (BIC) tests, as implemented in the jModelTest v0.1 software package [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. The Maximum Likelihood (ML) and Bayesian Inference (BI) were employed to infer phylogenetic relationships among genotypes and to construct phylogenetic trees. The ML analyses were performed using PhyML 3.0 [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e], under the evolutionary models selected according to the AIC and BIC criteria. The robustness of the phylogenetic relationships inferred from the ML trees were assessed using bootstrap analysis [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e], with 1,000 replicates. The Bayesian Inference (BI) analyses were performed using MrBayes v. 3.2 [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. The analyses were conducted for ten million generations (ngen\u0026thinsp;=\u0026thinsp;2,000,000) using two independent runs, each comprising four simultaneous Markov Chain Monte Carlo (MCMC) chains (one cold and three heated chains; nchains\u0026thinsp;=\u0026thinsp;4). Trees and model parameters were sampled every 1,000 generations (samplefreq\u0026thinsp;=\u0026thinsp;1000). The first 25% of samples were discarded as burn-in (burninfrac\u0026thinsp;=\u0026thinsp;0.25). The substitution model was set to GTR with a proportion of invariable sites and gamma-distributed rate variation among sites (lset nst\u0026thinsp;=\u0026thinsp;6, rates\u0026thinsp;=\u0026thinsp;invgamma). Nucleotide sequence similarities between genotypes were calculated using BioEdit v 7.2.5. The novel SSU rDNA genotypes obtained in this study were deposited in GenBank under accession numbers XXXXXXXX-XXXXXXXX.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eIn this study, a total of 330 specimens of \u003cem\u003eMullus barbatus\u003c/em\u003e L., 1758, 253 specimens of \u003cem\u003eMullus surmuletus\u003c/em\u003e L., 1758, and 240 specimens of \u003cem\u003eUpeneus moluccensis\u003c/em\u003e Bleeker, 1885, collected from three different locations (Muğla, Antalya, and Mersin) along the Mediterranean coast of T\u0026uuml;rkiye, were examined for myxosporean infections. As a result, three different myxosporean species belonging to three distinct genera were identified in the infected \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e specimens. However, no myxosporean infections were detected in any of the \u003cem\u003eUpeneus moluccensis\u003c/em\u003e specimens examined. The taxonomic summaries, morphological characteristics, and infection indices of the myxosporean species identified in this study are presented below;\u003c/p\u003e \u003cp\u003e \u003cb\u003eKudoa mediterraneus\u003c/b\u003e \u003cb\u003en. sp.\u003c/b\u003e\u003c/p\u003e\n\u003ch3\u003eTaxonomic summary\u003c/h3\u003e\n\u003cp\u003e \u003cstrong\u003ePhylum\u003c/strong\u003e \u003cp\u003eCnidaria Hatschek, 1888\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eClass\u003c/strong\u003e \u003cp\u003eMyxozoa Grasse, 1970\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSubclass\u003c/strong\u003e \u003cp\u003eMyxosporea B\u0026uuml;tschli, 1881\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eOrder\u003c/strong\u003e \u003cp\u003eMultivalvulida Shulman, 1959\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eFamily\u003c/strong\u003e \u003cp\u003eKudoidae Meglitsch, 1960\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eGenus\u003c/strong\u003e \u003cp\u003e \u003cem\u003eKudoa\u003c/em\u003e Meglitsch, 1947\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecies name\u003c/strong\u003e \u003cp\u003e \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType host\u003c/strong\u003e \u003cp\u003e \u003cem\u003eMullus barbatus\u003c/em\u003e L., 1758 (Red mullet) and \u003cem\u003eMullus surmelatus\u003c/em\u003e L., 1758 (Striped red mullet)\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType locality\u003c/strong\u003e \u003cp\u003eCoasts of Muğla (37\u0026deg; 12\u0026prime; 55\u0026Prime; N, 28\u0026deg; 21\u0026prime; 49\u0026Prime; E), Antalya (36\u0026deg; 14\u0026rsquo; 38\u0026rsquo;\u0026rsquo; N, 29\u0026deg; 59\u0026rsquo; 8\u0026rsquo;\u0026rsquo; E) and Mersin (36\u0026deg; 48\u0026prime; 44\u0026Prime; N, 34\u0026deg; 38\u0026prime; 29\u0026Prime; E), Mediterranean, T\u0026uuml;rkiye.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSite of infection\u003c/strong\u003e \u003cp\u003eMusculature.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType material\u003c/strong\u003e \u003cp\u003eOne holotype (E-5) and one paratype (E-12), fixed in 70% ethanol, were deposited in the Parasitological Collection of the Fatsa Faculty of Marine Sciences, Ordu University, Ordu-T\u0026uuml;rkiye.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNucleotide Sequences\u003c/strong\u003e \u003cp\u003eThe 18S rDNA nucleotide sequence obtained in this study was deposited in GenBank under accession number XXXXXXXX (specimen E-5).\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eEtymology\u003c/strong\u003e \u003cp\u003eThe specific epithet \u0026lsquo;\u003cem\u003emediterraneus\u003c/em\u003e\u0026rsquo; refers to the geographical region from which the species was described.\u003c/p\u003e \u003c/p\u003e\n\u003ch3\u003eDescription of myxospores\u003c/h3\u003e\n\u003cp\u003eNo plasmodium was observed. In apical view, the myxospores were somewhat stellate in shape, composed of four equal shell valves, with indistinct suture lines. The posterior poles were slightly angular. Four equal-sized, ovoid to drop-shaped polar capsules were present; however, the coils of the polar flaments were not visible (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea, c). In lateral view, the myxospores were typically flattened to slightly convex at the posterior pole, with a rounded anterior pole. The polar capsules are clearly teardrop-shaped (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Myxospores have the following dimensions (n\u0026thinsp;=\u0026thinsp;30): spore body length 5.35 (6.79\u0026ndash;4.12) \u0026micro;m, spore body width 8.24 (9.75\u0026ndash;7.19) \u0026micro;m, thickness 5.98 (6.96\u0026ndash;4.82) \u0026micro;m, polar capsule length 2.07 (2.99\u0026ndash;1.38) \u0026micro;m, polar capsule width 1.35 (1.94\u0026ndash;1.01) \u0026micro;m (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eMolecular phylogenetic analyses\u003c/h2\u003e \u003cp\u003eAs a result of nucleotide sequencing, 1319 bp of the 18S rDNA was obtained from the \u003cem\u003eKudoa\u003c/em\u003e specimen (E-5) selected for phylogenetic analyses. BLAST analysis indicated that this genotype was closely related to \u003cem\u003eKudoa unicapsula\u003c/em\u003e (AM490334; [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]), \u003cem\u003eK. parvibulbosa\u003c/em\u003e (LC626079; [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]), \u003cem\u003eK. trifolia\u003c/em\u003e (AM183300; [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]), and \u003cem\u003eK. iidae\u003c/em\u003e (LC493822; [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]). Subsequently, a dataset was constructed that included these species together with members of closely related (sister) sub-lineages within the genus \u003cem\u003eKudoa\u003c/em\u003e (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). The phylogenetic analyses were conducted over 1221 aligned nucleotide positions with 164 polymorphic sites (184 substitutional mutations). The AIC and BIC tests suggested the GTR\u0026thinsp;+\u0026thinsp;I+G (I:0.727, G: 0.685) and TPM3uf\u0026thinsp;+\u0026thinsp;I+G (I:0.738, G: 0.709) evolutionary models, respectively. The ML trees constructed under both models revealed similar topologies; however, the tree generated using the former model was supported by higher bootstrap values. The phylogenetic relationships inferred from the BI analysis were largely consistent with those obtained using the ML approach, with only minor differences that did not influence the phylogenetic placement of the newly identified genotype in the present study. In this study, the ML tree generated under the GTR\u0026thinsp;+\u0026thinsp;I+G model is presented due to its higher bootstrap support. Bootstrap values from the ML (GTR\u0026thinsp;+\u0026thinsp;I+G) analysis and posterior probability values from the BI analysis are shown on the tree (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In both phylogenetic trees generated using the ML and BI algorithms, the genotype obtained in this study (E-5) formed a lineage with \u003cem\u003eK. trifolia\u003c/em\u003e, \u003cem\u003eK. unicapsula\u003c/em\u003e and \u003cem\u003eK. iidae\u003c/em\u003e, showing nucleotide sequence similarities of 99.1%, 99.1% and 98.8%, respectively. In the ML tree, however, several additional species, including \u003cem\u003eK. parvibulbosa\u003c/em\u003e, \u003cem\u003eK. aburakarae\u003c/em\u003e and \u003cem\u003eK. javaensis\u003c/em\u003e, also placed in this lineage; nevertheless, their inclusion was not supported by significant bootstrap values (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In contrast, in the BI tree these three species formed a distinct lineage together with \u003cem\u003eK. quadricornis\u003c/em\u003e and \u003cem\u003eK. paraquadricornis\u003c/em\u003e, which was recovered as sister to the lineage containing the newly obtained genotype. In both cases, the phylogenetic relationships of the newly obtained genotype and \u003cem\u003eK. trifolia\u003c/em\u003e, \u003cem\u003eK. unicapsularis\u003c/em\u003e and \u003cem\u003eK. iidae\u003c/em\u003e were consistent and supported with sufficient bootstrap and posterior probability values.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eRemarks\u003c/h3\u003e\n\u003cp\u003eWhen the spores of \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. were compared with those of the phylogenetically closest species (\u003cem\u003eK. trifolia\u003c/em\u003e, \u003cem\u003eK. unicapsularis\u003c/em\u003e, \u003cem\u003eK. iidae\u003c/em\u003e, and \u003cem\u003eK. unicapsula\u003c/em\u003e) and the geographically closest species (\u003cem\u003eK. nil\u0026uuml;feri\u003c/em\u003e, \u003cem\u003eK. anatolica\u003c/em\u003e, and \u003cem\u003eKudoa dicentrarchi\u003c/em\u003e), all of these species were found to share unequal-sized valves and polar capsules. In contrast, \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. possesses four equal-sized valves and polar capsules. In particular, \u003cem\u003eK. unicapsularis\u003c/em\u003e, \u003cem\u003eK. iidae\u003c/em\u003e, and \u003cem\u003eK. unicapsula\u003c/em\u003e show pronounced variations in spore morphology that clearly distinguish them from \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. Among these species, \u003cem\u003eK. iidae\u003c/em\u003e exhibits a tail-like tapering extension of one of the shell valves, \u003cem\u003eK. unicapsula\u003c/em\u003e possesses one polar capsule that is significantly larger than the others, and \u003cem\u003eK. trifolia\u003c/em\u003e exhibits three small and one extremely large shell valves. Therefore, these species are morphologically very different from \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp and are unlikely to be confused with it. The geographically related species, on the other hand, all have genetic data (18S rDNA sequences) available in GenBank, which are clearly distinct from those of \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp.\u003c/p\u003e \u003cp\u003e \u003cb\u003ePrevalence, infection site(s), infection time and geographical distribution of\u003c/b\u003e \u003cb\u003eKudoa mediterraneus\u003c/b\u003e \u003cb\u003en. sp.\u003c/b\u003e\u003c/p\u003e \u003cp\u003eOut of 330 \u003cem\u003eMullus barbatus\u003c/em\u003e specimens, 16 were infected with \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp., corresponding to a prevalence of 4.9%. Additionally, this species was detected in one of 253 specimens of \u003cem\u003eM. surmelatus\u003c/em\u003e examined, representing a prevalence of 0.4%. However, no infection was detected in any of the 240 \u003cem\u003eUpeneus moluccensis\u003c/em\u003e specimens examined. All \u003cem\u003eK. mediterraneus\u003c/em\u003e specimens were observed in the muscles of the host fishes, and no clinical signs were detected in the infected tissues. Fifteen of the sixteen \u003cem\u003eK. mediterraneus\u003c/em\u003e specimens were detected in summer, whereas only one was found in autumn. Geographically, twelve of the sixteen \u003cem\u003eK. mediterraneus\u003c/em\u003e specimens were detected from Mersin (the eastern Mediterranean coasts of Turkiye), and four were detected in Antalya (the central Mediterranean coasts of Turkiye). However, no \u003cem\u003eK. mediterraneus\u003c/em\u003e infection was observed in Muğla (the western Mediterranean coasts of Turkiye).\u003c/p\u003e \u003cp\u003e \u003cb\u003eMyxobolus mullus\u003c/b\u003e \u003cb\u003en. sp.\u003c/b\u003e\u003c/p\u003e\n\u003ch3\u003eTaxonomic summary\u003c/h3\u003e\n\u003cp\u003e \u003cstrong\u003ePhylum\u003c/strong\u003e \u003cp\u003eCnidaria Hatschek, 1888\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eClass\u003c/strong\u003e \u003cp\u003eMyxozoa Grasse, 1970\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSubclass\u003c/strong\u003e \u003cp\u003eMyxosporea B\u0026uuml;tschli, 1881\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eOrder\u003c/strong\u003e \u003cp\u003eBivalvulida Shul\u0026rsquo;man, 1959\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eFamily\u003c/strong\u003e \u003cp\u003eMyxobolidae Th\u0026eacute;lohan, 1892\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eGenus\u003c/strong\u003e \u003cp\u003eMyxobolus B\u0026uuml;tschli, 1882\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecies name\u003c/strong\u003e \u003cp\u003e \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType host\u003c/strong\u003e \u003cp\u003e \u003cem\u003eMullus barbatus\u003c/em\u003e L., 1758 (Red mullet) and \u003cem\u003eMullus surmelatus\u003c/em\u003e L., 1758 (Striped red mullet)\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType locality\u003c/strong\u003e \u003cp\u003eCoasts of Muğla (37\u0026deg; 12\u0026prime; 55\u0026Prime; N, 28\u0026deg; 21\u0026prime; 49\u0026Prime; E), Antalya (36\u0026deg; 14\u0026rsquo; 38\u0026rsquo;\u0026rsquo; N, 29\u0026deg; 59\u0026rsquo; 8\u0026rsquo;\u0026rsquo; E) and Mersin (36\u0026deg; 48\u0026prime; 44\u0026Prime; N, 34\u0026deg; 38\u0026prime; 29\u0026Prime; E), Mediterranean, T\u0026uuml;rkiye.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSite of infection\u003c/strong\u003e \u003cp\u003eGall bladder\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType material\u003c/strong\u003e \u003cp\u003eHolotype (E-2) and one paratype (E-17), fixed in 70% ethanol, were deposited in the Parasitological Collection of the Fatsa Faculty of Marine Sciences, Ordu University, Ordu-T\u0026uuml;rkiye.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNucleotide Sequences\u003c/strong\u003e \u003cp\u003eThe 18S rDNA nucleotide sequence obtained in this study was deposited in GenBank under accession number XXXXXXXX (specimen E-2).\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eEtymology\u003c/strong\u003e \u003cp\u003eThe specific epithet \u0026lsquo;\u003cem\u003emullus\u003c/em\u003e\u0026rsquo; refers to the host fish genus from which this parasite species was described.\u003c/p\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eDescription of vegetative stage and myxospores\u003c/h2\u003e \u003cp\u003eNo plasmodia were observed. Fresh myxospores detected in the gall bladders were oval to oblong in valvular view and elliptical in sutural and apical views (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-f). The measurements of the spore body were as follows (n\u0026thinsp;=\u0026thinsp;30); 9.99 (11.11\u0026ndash;9.05) \u0026micro;m in length, 7.23 (7.97\u0026ndash;6.71) \u0026micro;m in width and 5.98 (6.53\u0026ndash;5.46) \u0026micro;m in thickness. The two slightly unequally sized polar capsules were pyriform in shape and positioned parallel to each other. The larger polar capsule measured 3.87 (4.46\u0026ndash;3.16) \u0026micro;m in length, 2.54 (2.91\u0026ndash;2.13) \u0026micro;m in width, whereas the smaller polar capsule measured 3.53 (4.19-3.0) \u0026micro;m in length, 2.36 (2.84\u0026ndash;2.02) \u0026micro;m in width (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSite of infection, hosts, geographical localities, and spore dimensions (\u0026micro;m) of \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp. identified in the present study, together with those of related myxozoan 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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSpore body\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePolar capsule\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSite of infection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eHost species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eReferences\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ethickness\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.99 (11.11\u0026ndash;9.05)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.23 (7.97\u0026ndash;6.71)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.98 (6.53\u0026ndash;5.46)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 3.87 (4.46\u0026ndash;3.16) Ls: 3.53 (4.19-3.00)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 2.54 (2.91\u0026ndash;2.13) Ls: 2.36 (2.84\u0026ndash;2.02)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eGall Bladder\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMullus Barbatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTurkish Coast of Mediterranean\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e(This Study)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus iwagiensis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e12.1 (12.8\u0026ndash;11.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.8 (10.5\u0026ndash;9.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.9 (8.2\u0026ndash;7.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 6.4 (6.9\u0026ndash;5.9)\u003c/p\u003e \u003cp\u003eLs: 5.8 (6.5\u0026ndash;5.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 4.0 (4.4\u0026ndash;3.4)\u003c/p\u003e \u003cp\u003eLs: 3.5 (4.0-3.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eBrain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eOryzias latipes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus tai\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10.2 (10.8\u0026ndash;9.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e7.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 (8.1\u0026ndash;7.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e5.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 (6.0-5.5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLp: 4.4 (4.9\u0026ndash;3.8)\u003c/p\u003e \u003cp\u003eLs: 4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 (4.5\u0026ndash;3.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLp: 2.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 (3.0-2.4)\u003c/p\u003e \u003cp\u003eLs: 2.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 (2.9\u0026ndash;2.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eBrain\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003ePagrus major\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eJapan\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u0026ndash;11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e6.5-7\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\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKidney connective tissue\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eSymphodus tinca\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eItaly\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e8\u0026ndash;10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.8\u0026ndash;6.5\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\u003e4.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKidney\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003eLabridae sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eUkraine\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e8.9\u0026ndash;10.8 (10.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.6\u0026ndash;6.9 (6.5)\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\u003e4.4\u0026ndash;5.3 (4.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e4.4\u0026ndash;5.3 (4.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKidney\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eParablennius tentacularis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTurkey: Sinop (Black Sea)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.0\u0026ndash;10.7 (7.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e5.1\u0026ndash;7.0 (4.3)\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\u003e4.5\u0026ndash;5.3 (5.0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.9\u0026ndash;3.4 (3.1)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eKidney\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eParablennius sanguinolentus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eTurkey: Sinop (Black Sea)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]\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\u003eMolecular phylogenetic analyses\u003c/h2\u003e \u003cp\u003eA 1585 bp fragment of the 18S rDNA gene was obtained from the \u003cem\u003eMyxobolus\u003c/em\u003e sp. specimen (E-2) selected for phylogenetic analyses. The 18S rDNA genotypes of two recently described species, \u003cem\u003eMyxobolus iwagiensis\u003c/em\u003e Kawano, Sakurai \u0026amp; Yanagida, 2025 (LC861743; [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]) and \u003cem\u003eMyxobolus tai\u003c/em\u003e Kawano, Nitta \u0026amp; Yanagida, 2026 (LC884833, LC886104; [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]), were identified in BLAST analysis as the most similar sequences to the genotype newly obtained. Accordingly, a dataset comprising these two species and other closely related myxozoan species was constructed for further phylogenetic analyses (Table S2). However, the dataset was limited to 889 bp, including 436 variable sites and 659 mutations, due to the short length of some sequences and the high number of insertion and deletion mutations. The GTR\u0026thinsp;+\u0026thinsp;I+G (I:0.231, G: 0.592) and TIM2\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G (I:0.228, G: 0.578) evolutionary models were selected based on the AIC and BIC tests, respectively. The ML trees constructed under both models showed similar topologies, although higher bootstrap values were obtained under the first model. The BI analysis, conducted under GTR\u0026thinsp;+\u0026thinsp;I+G mode, produced a tree with a topology similar to that of the ML tree, showing only minor variation in the interlineage relationships within the first lineage. Due to its higher bootstrap support, the ML tree constructed with the GTR\u0026thinsp;+\u0026thinsp;I+G model is presented in Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. Additionally, bootstrap values from the ML (GTR\u0026thinsp;+\u0026thinsp;I+G) analysis and posterior probability values from the BI analysis are indicated on the tree. In the phylogenetic trees generated using the ML and BI algorithms, three similar lineages were observed, one of which (the third lineage) comprised the newly obtained genotype in this study (E-2), together with \u003cem\u003eMyxobolus tai\u003c/em\u003e and \u003cem\u003eMyxobolus iwagiensis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). On both trees, the newly obtained genotype in this lineage appeared as sister taxa to \u003cem\u003eM. tai\u003c/em\u003e genotypes, and this relationship was supported by 68% bootstrap value and 0.92 posterior probability. The nucleotide sequence similarity between E-2 and \u003cem\u003eM. tai\u003c/em\u003e genotypes were 94.4%. The third species of the lineage, \u003cem\u003eM. iwagiensis\u003c/em\u003e, placed as sister taxon to the previously mentioned species with 100% bootstrap support and 1.0 posterior probability. Additionally, the nucleotide sequence similarity between E-2 and \u003cem\u003eM. iwagiensis\u003c/em\u003e genotype was 92.7%.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eRemarks\u003c/h2\u003e \u003cp\u003eUp to date, a total of nine \u003cem\u003eMyxobolus\u003c/em\u003e species have been reported from the Turkish coasts, four of which possess unequal-sized polar capsules similar to those observed in the specimens obtained in the present study. However, three of these species have available genetic data (18S rDNA sequences) in GenBank that do not match the genotypes obtained in this study. One species, \u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e Parasi, 1912, lacks genetic data and therefore can only be compared based on morphological characteristics. When the spore measurements of \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp. are compared with those of \u003cem\u003eM. asymmetricus\u003c/em\u003e and the two phylogenetically closest species, \u003cem\u003eMyxobolus iwagiensis\u003c/em\u003e and \u003cem\u003eMyxobolus tai\u003c/em\u003e, both the spore body and polar capsule measurements are significantly smaller than those of \u003cem\u003eM. iwagiensis\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]. Furthermore, the two polar capsules of \u003cem\u003eM. iwagiensis\u003c/em\u003e were markedly unequal in size, whereas those of \u003cem\u003eM. mullus\u003c/em\u003e n. sp. were only slightly unequal. Although the spore body measurements of \u003cem\u003eM. tai\u003c/em\u003e and \u003cem\u003eM. mullus\u003c/em\u003e n. sp. are quite similar, the polar capsules of \u003cem\u003eM. tai\u003c/em\u003e are considerably longer (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Finally, while the spore body lengths of \u003cem\u003eM. asymmetricus\u003c/em\u003e and \u003cem\u003eM. mullus\u003c/em\u003e n. sp. are quite similar, the spore body width is significantly greater in the \u003cem\u003eM. mullus\u003c/em\u003e n. sp. (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cb\u003ePrevalence, infection site(s), infection time and geographical distribution of\u003c/b\u003e \u003cb\u003eMyxobolus mullus\u003c/b\u003e \u003cb\u003en. sp.\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp. was detected in 23 of 330 \u003cem\u003eMullus barbatus\u003c/em\u003e specimens, corresponding to a prevalence of 7.0%. Additionally, the species was observed in 27 of 253 \u003cem\u003eM. surmelatus\u003c/em\u003e specimens, corresponding to a prevalence of 10.7%. However, no infection with \u003cem\u003eM. mullus\u003c/em\u003e n. sp. was detected in \u003cem\u003eUpeneus moluccensis\u003c/em\u003e. All specimens of \u003cem\u003eM. mullus\u003c/em\u003e n. sp. observed in both \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmelatus\u003c/em\u003e were recovered from the gall bladder. However, no clinical signs were observed in any the infected host fishes. Seasonally, majority of the \u003cem\u003eM. mullus\u003c/em\u003e n. sp. infections (\u003cem\u003en\u003c/em\u003e: 44) were detected during the spring and summer periods, whereas a few infections (\u003cem\u003en\u003c/em\u003e: 6) were recorded in the winter and autumn periods. Geographically, of the 50 \u003cem\u003eM. mullus\u003c/em\u003e n. sp. specimens, 35 were detected from Mersin (the eastern Mediterranean coasts of Turkiye), 10 were detected in Antalya (the central Mediterranean coasts of Turkiye), and 5 were detected in Muğla (the western Mediterranean coasts of Turkiye).\u003c/p\u003e \u003cp\u003e \u003cb\u003eOrtholinea mullusi\u003c/b\u003e \u003cb\u003eG\u0026uuml;rkanlı, Okkay, \u0026Ccedil;ift\u0026ccedil;i, Yurakhno and \u0026Ouml;zer 2018.\u003c/b\u003e\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eTaxonomic summary\u003c/h2\u003e \u003cp\u003e \u003cstrong\u003ePhylum\u003c/strong\u003e \u003cp\u003eCnidaria Hatschek, 1888\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eClass\u003c/strong\u003e \u003cp\u003eMyxozoa Grass\u0026eacute;, 1970\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSubclass\u003c/strong\u003e \u003cp\u003eMyxosporea B\u0026uuml;tschli, 1881\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eOrder\u003c/strong\u003e \u003cp\u003eBivalvulida Shul\u0026rsquo;man, 1959\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eFamily\u003c/strong\u003e \u003cp\u003eOrtholineidae Lom \u0026amp; Noble, 1984\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eGenus\u003c/strong\u003e \u003cp\u003e \u003cem\u003eOrtholinea\u003c/em\u003e Shul\u0026rsquo;man, 1962\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSpecies name\u003c/strong\u003e \u003cp\u003e \u003cem\u003eOrtholinea mullusi\u003c/em\u003e G\u0026uuml;rkanlı, Okkay, \u0026Ccedil;ift\u0026ccedil;i, Yurakhno \u0026amp; \u0026Ouml;zer, 2018\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType host\u003c/strong\u003e \u003cp\u003e \u003cem\u003eMullus barbatus\u003c/em\u003e L., 1758\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eType locality\u003c/strong\u003e \u003cp\u003eCoast of Sinop, Black Sea, T\u0026uuml;rkiye (42\u0026deg; 02\u0026rsquo; 51\u0026rsquo;\u0026rsquo; N, 35\u0026deg; 02\u0026rsquo; 56\u0026rsquo;\u0026rsquo; E).\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNew locality\u003c/strong\u003e \u003cp\u003eCoasts of Muğla (37\u0026deg; 12\u0026prime; 55\u0026Prime; N, 28\u0026deg; 21\u0026prime; 49\u0026Prime; E) and Antalya (36\u0026deg; 14\u0026rsquo; 38\u0026rsquo;\u0026rsquo; N, 29\u0026deg; 59\u0026rsquo; 8\u0026rsquo;\u0026rsquo; E), Mediterranean, T\u0026uuml;rkiye.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eSite of infection\u003c/strong\u003e \u003cp\u003eUrinary bladder and kidney tubules.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNew site(s) of infection\u003c/strong\u003e \u003cp\u003eGonads, Kidney and Urinary bladder\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eNucleotide Sequences\u003c/strong\u003e \u003cp\u003eThe 18S rDNA nucleotide sequences obtained in this study were deposited in GenBank under accession numbers XXXXXXXX (specimen E-7), XXXXXXXX (specimen E-9) and XXXXXXXX (specimen E-13).\u003c/p\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eDescription of myxospores\u003c/h2\u003e \u003cp\u003eNo plasmodium was observed. In frontal view, mature spores were spherical with rounded anterior and posterior poles (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea). However, in sutural and apical views, the spores were ellipsoidal (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003eb, c). Two polar capsules were equal in size and somewhat spherical. Notably, no polar flaments were observed. The dimensions of myxospores (n\u0026thinsp;=\u0026thinsp;30 from each host) were as follows: spore body length 9.16 (9.52\u0026ndash;8.92), width 8.33 (9.16\u0026ndash;7.83), polar capsul length 3.16 (3.25\u0026ndash;2.93) and polar capsul width 2.50 (2.62\u0026ndash;2.41) (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\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\u003eSite of infection, hosts, geographical localities, and spore dimensions (\u0026micro;m) of \u003cem\u003eOrtholinea mullusi\u003c/em\u003e identified in the present study, together with those of related myxozoan 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\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eSpore body\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ePolar capsule\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSite of infection\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eHost species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eLocality\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eReferences\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ethickness\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003elength\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003ewidth\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eOrtholinea mullusi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.16\u003c/p\u003e \u003cp\u003e(8.92\u0026ndash;9.52)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.33\u003c/p\u003e \u003cp\u003e(7.83\u0026ndash;9.16)\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\u003e3.16\u003c/p\u003e \u003cp\u003e(2.93\u0026ndash;3.25)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.50\u003c/p\u003e \u003cp\u003e(2.41\u0026ndash;2.62)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eGonads, kidney, urinary bladder\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMullus barbatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eMuğla and Antalya coasts of the Mediterranean, T\u0026uuml;rkiye\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e(This study)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eOrtholinea mullusi\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.3\u003c/p\u003e \u003cp\u003e(9.0\u0026thinsp;\u0026minus;\u0026thinsp;9.7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.7\u003c/p\u003e \u003cp\u003e(8.2\u0026thinsp;\u0026minus;\u0026thinsp;9.3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.7\u003c/p\u003e \u003cp\u003e(7.5\u0026thinsp;\u0026minus;\u0026thinsp;7.9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.1\u003c/p\u003e \u003cp\u003e(3.0-3.2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.5\u003c/p\u003e \u003cp\u003e(2.4\u0026ndash;2.6)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003eUrinary bladder, kidney\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e\u003cem\u003eMullus barbatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003eSinop coasts of the Black Sea, T\u0026uuml;rkiye\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c10\"\u003e \u003cp\u003e[\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]\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=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eMolecular phylogenetic analyses\u003c/h2\u003e \u003cp\u003eNucleotide sequencings yielded approximately 850\u0026ndash;1700 bp of the 18S rDNA gene from the three \u003cem\u003eOrtholinea\u003c/em\u003e specimens (E-7, E-9 and E-13), which were subsequently subjected to phylogenetic analyses. All three specimens exhibited identical 18S rDNA sequences. BLAST analyses revealed \u003cem\u003eOrtholinea mullusi\u003c/em\u003e (MF539825; [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]) as the closest species to all newly obtained \u003cem\u003eOrtholinea\u003c/em\u003e specimens (Table S3). For phylogenetic analyses, a dataset including \u003cem\u003eO. mullusi\u003c/em\u003e and related \u003cem\u003eOrtholinea\u003c/em\u003e species was constructed. The final alignment comprised 832 nucleotide positions, of which 281 were polymorphic (409 substitutional mutations). The AIC and BIC tests suggested GTR\u0026thinsp;+\u0026thinsp;I+G (I:0.39, G: 0.49) and TPM2uf\u0026thinsp;+\u0026thinsp;I+G (I:0.379, G: 0.46), evolutionary models, respectively. Similar phylogenetic relationships were obtained in the ML analyses conducted under both models. Likewise, the BI analysis produced a tree topology largely consistent with those inferred from the ML analysis. The ML tree generated under the GTR\u0026thinsp;+\u0026thinsp;I+G model is presented here due to its higher bootstrap support. Posterior probability values from the BI analysis are shown at the corresponding nodes together with ML bootstrap values (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). In both ML and BI trees, the newly obtained \u003cem\u003eOrtholinea\u003c/em\u003e genotype (E-7) clustered with \u003cem\u003eO. mullusi\u003c/em\u003e, showing 99.3% nucleotide sequence similarity. This relationship was strongly supported by 97% bootstrap and a posterior probability of 1.0. \u003cem\u003eOrtholinea\u003c/em\u003e sp. specimen RT-1 (MK937851; [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]) appeared as a sister to this lineage, with 54% bootstrap support and a posterior probability of 0.81.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eRemarks\u003c/h2\u003e \u003cp\u003eComparison of the three \u003cem\u003eOrtholinea\u003c/em\u003e specimens obtained in this study with \u003cem\u003eO. mullusi\u003c/em\u003e, their closest phylogenetic relative, revealed a high degree of morphological similarity of the myxospores in frontal (spherical), sutural (ellipsoidal), and apical (spherical) views (Fig.\u0026nbsp;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003ea-c). Likewise, the spor body length and width as well as the polar capsul length and width, of the newly obtaned \u003cem\u003eOrtholinea\u003c/em\u003e specimens were fully consistent with those of \u003cem\u003eO. mullusi\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The only difference between them was the shape of the polar capsules: in the original description of \u003cem\u003eO. mullusi\u003c/em\u003e, the polar capsules were pyriform and equal in size, whereas in the newly obtained specimens, they were also equal in size but nearly ovoidal in shape.\u003c/p\u003e \u003cp\u003e \u003cb\u003ePrevalence, infection site(s), infection time and geographical distribution of\u003c/b\u003e \u003cb\u003eOrtholinea mullusi\u003c/b\u003e\u003c/p\u003e \u003cp\u003eOnly three out of 330 examined \u003cem\u003eM. barbatus\u003c/em\u003e specimens were found to be infected with \u003cem\u003eO. mullusi\u003c/em\u003e, corresponding to a prevalence of 0.9%. However, no infection was detected in \u003cem\u003eM. surmelatus\u003c/em\u003e and \u003cem\u003eU. moluccensis\u003c/em\u003e specimens. All three \u003cem\u003eO. mullusi\u003c/em\u003e infections were observed in different host organs including, the gonads, urinary bladder and kidney. Additionally, no clinical signs were detected in the infected organs. Two of the three \u003cem\u003eO. mullusi\u003c/em\u003e infections were recorded in host specimens collected from Antalya (the central Mediterranean coasts of Turkiye) in summer, while one was detected in autumn from Muğla (the the western Mediterranean coasts of Turkiye).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn the Mediterranean Sea, the family Mullidae, which includes economically important species, comprises six species, five of which have also been reported from the Mediterranean coasts of T\u0026uuml;rkiye [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. Despite their economic importance, there are only a few records concerning the myxozoan diversity infecting members of the Mullidae family, and none of them relate to the western Mediterranean [\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e, \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e]. So, this study provided new results on the myxozoan diversity infecting \u003cem\u003eMullus barbatus\u003c/em\u003e, \u003cem\u003eMullus surmuletus\u003c/em\u003e, and \u003cem\u003eUpeneus moluccensis\u003c/em\u003e collected from three locations, Muğla, Antalya, and Mersin along the Mediterranean coasts of T\u0026uuml;rkiye.\u003c/p\u003e \u003cp\u003eInitial microscopic examinations revealed specimens belonging to three different myxozoan genera, \u003cem\u003eKudoa\u003c/em\u003e Meglitsch, 1947, \u003cem\u003eMyxobolus\u003c/em\u003e B\u0026uuml;tschli, 1882, and \u003cem\u003eOrtholinea\u003c/em\u003e Shulman, 1962, in various organs of \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e. However, no myxosporean infections were detected in \u003cem\u003eUpeneus moluccensis\u003c/em\u003e specimens. The \u003cem\u003eKudoa\u003c/em\u003e infections were exclusively detected in the trunk muscles of \u003cem\u003eM. barbatus\u003c/em\u003e (\u003cem\u003en\u003c/em\u003e: 16) and \u003cem\u003eM. surmuletus\u003c/em\u003e (\u003cem\u003en\u003c/em\u003e: 1) specimens. The complete similarity in myxospor morphology, the concordance of morphometric data, and the shared source tissue (trunk muscles) indicate that all specimens belonged to the same \u003cem\u003eKudoa\u003c/em\u003e species (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea-c; Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). In the phylogenetic trees constructed using the ML and BI algorithms, the \u003cem\u003eKudoa\u003c/em\u003e sp. genotype obtained in this study (E-5) formed a distinct lineage together with \u003cem\u003eKudoa trifoli\u003c/em\u003e, \u003cem\u003eKudoa unicapsula\u003c/em\u003e, and \u003cem\u003eKudoa iidae\u003c/em\u003e, supported by high bootstrap values and posterior probabilities (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Within the genus \u003cem\u003eKudoa\u003c/em\u003e, interspecific nucleotide sequence similarity based on 18S rDNA is generally high compared with other myxozoan genera. For example, the similarity is 99.9% between \u003cem\u003eKudoa quadricornis\u003c/em\u003e and \u003cem\u003eKudoa paraquadricornis\u003c/em\u003e, 99.8% between \u003cem\u003eKudoa yasunagai\u003c/em\u003e and \u003cem\u003eKudoa chaetodoni\u003c/em\u003e, and 99.3% between \u003cem\u003eKudoa grammatorcyni\u003c/em\u003e and \u003cem\u003eKudoa scomberomori\u003c/em\u003e [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. From this perspective, the nucleotide sequence similarities between the new genotype identified in the present study and the above-mentioned species (ranging from 99.1% to 98.8%) are not sufficiently high to assign the new genotype to any of these previously described species. This assumption was further supported by the myxospore morphology and morphometric data (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). All phylogenetically related species-\u003cem\u003eK. trifolia\u003c/em\u003e, \u003cem\u003eK. unicapsula\u003c/em\u003e, and \u003cem\u003eK. iidae\u003c/em\u003e-possess unequal shell valves and polar capsules [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. In contrast, the specimens obtained in the present study exhibited morphologically homogeneous spores with equally sized shell valves and polar capsules, clearly distinguishing them from the aforementioned species. Based on the morphological, morphometric, and molecular phylogenetic data, we propose that the newly obtained \u003cem\u003eKudoa\u003c/em\u003e specimens represent a new species, designated as \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp.\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eKudoa\u003c/em\u003e currently includes 134 binomial species. Of these, three species have so far been reported from the Turkish seas: \u003cem\u003eK. dicentrarchi\u003c/em\u003e Sitja-Bobadilla \u0026amp; Alvarez-Pellitera, 1992 (ex \u003cem\u003eDicentrarchus labrax\u003c/em\u003e), \u003cem\u003eK. anatolica\u003c/em\u003e \u0026Ouml;zer, Okkay, G\u0026uuml;rkanlı, Yılmaz, Yurakhno, 2018 (ex \u003cem\u003eAtherina hepsetus\u003c/em\u003e) and \u003cem\u003eK. nil\u0026uuml;feri\u003c/em\u003e \u0026Ouml;zer, Okkay, G\u0026uuml;rkanlı, Yılmaz, Yurakhno, 2018 (ex \u003cem\u003eNeogobius melanostomus\u003c/em\u003e) [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e]. Accordingly, \u003cem\u003eK. mediterraneus\u003c/em\u003e n. sp. represents the forth species of \u003cem\u003eKudoa\u003c/em\u003e recorded from the Turkish coasts.\u003c/p\u003e \u003cp\u003eSo far, species of the genus \u003cem\u003eKudoa\u003c/em\u003e have been reported to infect a wide variety of fish species across different families [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e]. However, to the best of our knowledge, there has been only one record of a \u003cem\u003eKudoa\u003c/em\u003e species, \u003cem\u003eK. lutjanus\u003c/em\u003e Wang, Huang, Tsai, Cheng, Tsai, Chen, Chen, Chu, Liaw, Chang \u0026amp; Chen, 2005, infecting a member of the Mullidae, specifically \u003cem\u003eUpeneus tragula\u003c/em\u003e in China [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. Therefore, \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp. is the second \u003cem\u003eKudoa\u003c/em\u003e species described from a host in the family Mullidae worldwide. Additionally, it also represents the first \u003cem\u003eKudoa\u003c/em\u003e species known to infect mullid hosts in the Mediterranean Sea.\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eKudoa\u003c/em\u003e, which forms the histozoic lineage of Myxozoa, has been mostly found in the skeletal muscle of fish hosts. Additionally, infections have also been reported in tissues of various other organs, including the gills, brain, heart, kidney, spleen, ovary, gall bladder, urinary bladder, oesophagus, intestine, mesentery, and smooth muscle [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. Consistent with this, all \u003cem\u003eKudoa mediterraneus\u003c/em\u003e specimens in this study were determined in the trunk muscles. Although some \u003cem\u003eKudoa\u003c/em\u003e species, such as \u003cem\u003eK. thyrsites\u003c/em\u003e, \u003cem\u003eK. musculoliquefaciens\u003c/em\u003e, and \u003cem\u003eK. clupeidae\u003c/em\u003e, have been linked to post-harvest softening of fish flesh, no such pathological effects have been observed for \u003cem\u003eK. mediterraneus\u003c/em\u003e [\u003cspan additionalcitationids=\"CR54\" citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e]. Species of the genus \u003cem\u003eKudoa\u003c/em\u003e have been reported to reach peak infection rates during summer and autumn [\u003cspan additionalcitationids=\"CR57\" citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e]. In full agreement with this pattern, sixteen of the seventeen \u003cem\u003eKudoa mediterraneus\u003c/em\u003e specimens were found in summer, with only one specimen observed in autumn. The prevalence of \u003cem\u003eK. mediterraneus\u003c/em\u003e was 4.9% (16/330) in \u003cem\u003eM. barbatus\u003c/em\u003e and 0.4% (1/253) in \u003cem\u003eM. surmuletus\u003c/em\u003e, while no infection was detected in the examined \u003cem\u003eU. moluccensis\u003c/em\u003e specimens examined. When compared to prevalences reported for other \u003cem\u003eKudoa\u003c/em\u003e species from the Mediterranean and adjacent seas (the Aegean Sea, Sea of Marmara, and Black Sea)\u0026mdash;including \u003cem\u003eK. unicapsula\u003c/em\u003e from \u003cem\u003eLiza ramada\u003c/em\u003e (40%) and \u003cem\u003eLiza aurata\u003c/em\u003e (15%), \u003cem\u003eK. camarguensis\u003c/em\u003e from \u003cem\u003ePomatoschistus minutus\u003c/em\u003e (2.02%) and \u003cem\u003ePomatoschistus microps\u003c/em\u003e (12.58%), \u003cem\u003eK. nil\u0026uuml;feri\u003c/em\u003e from \u003cem\u003eNeogobius melanostomus\u003c/em\u003e (12.8%), and \u003cem\u003eK. anatolica\u003c/em\u003e from \u003cem\u003eAtherina hepsetus\u003c/em\u003e (32.1%)\u0026mdash;the infection levels observed in the present study seem comparatively low [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e59\u003c/span\u003e]. Moreover, these values are much lower than the prevalence reported for \u003cem\u003eK. lutjanus\u003c/em\u003e, the only \u003cem\u003eKudoa\u003c/em\u003e species previously documented from a mullid host, in \u003cem\u003eU. tragula\u003c/em\u003e (20%) [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. However, considering the prevalence of \u003cem\u003eK. mediterraneus\u003c/em\u003e n. sp. in \u003cem\u003eM. barbatus\u003c/em\u003e separately by location, it was 0.8%, 1.7%, and 13.3% in Muğla (western Mediterranean coast of T\u0026uuml;rkiye), Antalya (central Mediterranean coast of T\u0026uuml;rkiye), and Mersin (eastern Mediterranean coast of T\u0026uuml;rkiye), respectively. Additionally, only one \u003cem\u003eK. mediterraneus\u003c/em\u003e specimen was found in \u003cem\u003eM. surmuletus\u003c/em\u003e during the summer period in Antalya (one of 120 specimens examined, corresponding to a prevalence of 0.8%). These patterns suggest that \u003cem\u003eM. barbatus\u003c/em\u003e might be the main host for this parasite, which could be more prevalent in the eastern Mediterranean and might still be spreading toward the western Mediterranean.\u003c/p\u003e \u003cp\u003eThe second and most abundant myxozoan specimens identified in this study belonged to the genus \u003cem\u003eMyxobolus\u003c/em\u003e B\u0026uuml;tschli, 1882, accounting for 50 infections among a total of 823 examined specimens of \u003cem\u003eM. barbatus\u003c/em\u003e, \u003cem\u003eM. surmuletus\u003c/em\u003e, and \u003cem\u003eU. moluccensis\u003c/em\u003e. All observed \u003cem\u003eMyxobolus\u003c/em\u003e specimens showed similar myxospore morphology and morphometric characteristics (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea-f; Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Additionally, all specimens were found exclusively in the same organ, the gall bladder. All these data indicate that the specimens belong to the same \u003cem\u003eMyxobolus\u003c/em\u003e species. In both ML and BI trees based on 18S rDNA nucleotide sequences, the newly obtained genotype (E-2) formed a lineage with \u003cem\u003eMyxobolus tai\u003c/em\u003e [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e] and \u003cem\u003eM. iwagiens\u003c/em\u003e [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e] that supported by significant bootstrap values and posterior probabilities (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Considering the 18S rDNA nucleotide sequence identities between some closely related \u003cem\u003eMyxobolus\u003c/em\u003e species in our dataset, these values range from 96.9% to 98.2% (Table S4), indicating that sequence identities below the minimum threshold of 98.2% may correspond to distinct species. This threshold value for species delineation may be even higher when all \u003cem\u003eMyxobolus\u003c/em\u003e species are considered. In the light of this data, the nucleotide sequence identities were insufficient to assign the specimens obtained in this study to either of \u003cem\u003eM. tai\u003c/em\u003e and \u003cem\u003eM. iwagiensis\u003c/em\u003e, as the sequence similarities between the new genotype (E-2) and \u003cem\u003eM. tai\u003c/em\u003e and \u003cem\u003eM. iwagiensis\u003c/em\u003e were 94.4% and 92.7%, respectively, which are well below the 98.2% threshold. The morphology and morphometric characteristics of the myxospores also supported the molecular findings. Although the newly obtained \u003cem\u003eMyxobolus\u003c/em\u003e specimens and the two phylogenetically closest species, \u003cem\u003eM. tai\u003c/em\u003e and \u003cem\u003eM. iwagiensis\u003c/em\u003e, share a similar oblong spore shape, \u003cem\u003eM. iwagiensis\u003c/em\u003e differs from the new specimens by having larger spore body dimensions, while \u003cem\u003eM. tai\u003c/em\u003e has larger polar capsules compared to those of the new specimens [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]. \u003cem\u003eMyxobolus asymmetricus\u003c/em\u003e (lacking molecular data), a species previously reported from T\u0026uuml;rkiye, shows significantly narrower polar capsules than the new specimens (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]. Finally, \u003cem\u003eM. parvus\u003c/em\u003e, reported from \u003cem\u003eM. barbatus\u003c/em\u003e in the Black Sea, differs from the new specimens by having equal-sized polar capsules [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. Based on the combined morphological, morphometric, and molecular phylogenetic data, the examined \u003cem\u003eMyxobolus\u003c/em\u003e specimens are described here as a new species, \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp.\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eMyxobolus\u003c/em\u003e B\u0026uuml;tschli, 1882 is the most speciouse group within the phylum Myxozoa, comprising nearly 1000 valid species [\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e]. Of these, nine species-\u003cem\u003eM. asymmetricus\u003c/em\u003e Lalitha-Kumari, 1969 (ex. \u003cem\u003eParablennius sanguinolentus\u003c/em\u003e and \u003cem\u003eParablennius tentacularis\u003c/em\u003e), \u003cem\u003eM. episquamalis\u003c/em\u003e Egusa, Maeno \u0026amp; Sorimachi (ex. \u003cem\u003eMugil cephalus\u003c/em\u003e), \u003cem\u003eM. exiguus\u003c/em\u003e (ex. \u003cem\u003eMugil cephalus\u003c/em\u003e), \u003cem\u003eM. ichkeulensis\u003c/em\u003e (ex. \u003cem\u003eMugil cephalus\u003c/em\u003e), \u003cem\u003eM. muelleri\u003c/em\u003e B\u0026uuml;tschli, 1882 (ex. \u003cem\u003eMugil cephalus\u003c/em\u003e and \u003cem\u003eDiplodus annularis\u003c/em\u003e), \u003cem\u003eM. parvum\u003c/em\u003e Yurakhno, 1991 (ex. \u003cem\u003eParablennius tentacularis\u003c/em\u003e), \u003cem\u003eM. parvus\u003c/em\u003e Shulman, 1962 (ex. \u003cem\u003eLiza saliens\u003c/em\u003e, \u003cem\u003eChelon saliens\u003c/em\u003e and \u003cem\u003eMullus barbatus\u003c/em\u003e), \u003cem\u003eM. rotundus\u003c/em\u003e Nemeczek, 1911 (ex. \u003cem\u003eSymphodus cinereus\u003c/em\u003e) and \u003cem\u003eM. spinacurvatura\u003c/em\u003e Maeno, Sorimachi, Ogawa \u0026amp; Egusa, 1990 (ex. \u003cem\u003eMugil cephalus\u003c/em\u003e)- have been reported from the Turkish coasts [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan additionalcitationids=\"CR63 CR64 CR65\" citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e]. Moreover, six of these species have been reported from the Black Sea coasts of T\u0026uuml;rkiye, while two were recorded from the Mediterranean Sea and one from the Sea of Marmara. Therefore, \u003cem\u003eM. mullus\u003c/em\u003e represents the tenth \u003cem\u003eMyxobolus\u003c/em\u003e species reported from the Turkish coasts and the third \u003cem\u003eMyxobolus\u003c/em\u003e species recorded from the Mediterranean coast of T\u0026uuml;rkiye. Moreover, as mentioned earlier, only a limited number of myxozoan species have been reported as parasites of the Mullidae family, one of which is \u003cem\u003eM. parvus\u003c/em\u003e, described from the Black Sea [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. Accordingly, \u003cem\u003eM. mullus\u003c/em\u003e represents the second \u003cem\u003eMyxobolus\u003c/em\u003e species reported from \u003cem\u003eM. barbatus\u003c/em\u003e worldwide.\u003c/p\u003e \u003cp\u003eBased on the infection characteristics of \u003cem\u003eM. mullus\u003c/em\u003e n. sp., this species reached its peak infection rate (44 out of 50 cases) during the spring and summer seasons, when sea temperatures range from approximately 18\u0026deg;C to 28\u0026deg;C [\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e]. This pattern aligns with previously reported infection periods of \u003cem\u003eMyxobolus\u003c/em\u003e species [\u003cspan additionalcitationids=\"CR69\" citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e]. All fifty \u003cem\u003eM. mullus\u003c/em\u003e specimens in this study were found exclusively in the gall bladder. However, the organ -or tissue- specificity of this species remains uncertain. Several \u003cem\u003eMyxobolus\u003c/em\u003e species reported from the gall bladder\u0026mdash;\u003cem\u003eM. bankimi\u003c/em\u003e Sarkar, 1999; \u003cem\u003eM. chilkensis\u003c/em\u003e Kalavati, Venkateswara Rao \u0026amp; Vaidahei, 1992; \u003cem\u003eM. myleus\u003c/em\u003e Azevedo, Clemente, Casal, Matos, Alves, Al-Quraishy \u0026amp; Matos, 2012; \u003cem\u003eM. hepatobiliaris\u003c/em\u003e Rocha, Casal, Alves, Antunes, Rodrigues \u0026amp; Azevedo, 2019; and \u003cem\u003eM. vesicularis\u003c/em\u003e Rocha, Casal, Alves, Antunes, Rodrigues \u0026amp; Azevedo, 2019\u0026mdash;have not been reported from other organs or tissues so far. Conversely, some species, such as \u003cem\u003eM. cuneus\u003c/em\u003e Adriano, Arana \u0026amp; Cordeiro, 2006 and \u003cem\u003eM. parvus\u003c/em\u003e, have been documented from various organs -including the urinary bladder, gills, spleen, fins, head surface, lower jaw, liver, heart, kidney tubules, and within several cysts- in addition to the gall bladder [\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e, \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e71\u003c/span\u003e]. Therefore, further research is necessary to clarify this issue. Overall, the prevalence of \u003cem\u003eM. mullus\u003c/em\u003e n. sp. in \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e was 7% and 10.7%, respectively. These rates appear relatively low compared with previous reports of various \u003cem\u003eMyxobolus\u003c/em\u003e species recorded in T\u0026uuml;rkiye, where prevalence values ranged from 9% to 53% (general around 20\u0026ndash;25% for most species) [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan additionalcitationids=\"CR63 CR64 CR65\" citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e]. However, when analyzed by sampling location, the prevalence of \u003cem\u003eM. mullus\u003c/em\u003e n. sp. in Muğla (the western Mediterranean coast of T\u0026uuml;rkiye) was 12.5% in \u003cem\u003eM. barbatus\u003c/em\u003e and 27.4% in \u003cem\u003eM. surmuletus\u003c/em\u003e. In contrast, prevalence was only 3.3% and 5% in Antalya (central Mediterranean coast of T\u0026uuml;rkiye) for \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e, respectively, and 4.4% and 1.7% in Mersin (eastern Mediterranean coast of T\u0026uuml;rkiye) (Table S5). In conclusion, 70% (35 out of 50) of all \u003cem\u003eM. mullus\u003c/em\u003e n. sp. infections identified in this study were recorded from the western Mediterranean region. If the sea surface temperature factor is disregarded, since temperatures were similar across the sampling locations during the same months as mentioned in Şişman et al. [\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e], this pattern suggests that \u003cem\u003eM. mullus\u003c/em\u003e n. sp. may still be spreading from the western Mediterranean to the eastern Mediterranean.\u003c/p\u003e \u003cp\u003eMicroscopic examination of three \u003cem\u003eM. barbatus\u003c/em\u003e specimens (out of 330 examined) revealed a third myxozoan parasite belonging to the genus \u003cem\u003eOrtholinea\u003c/em\u003e Shulman, 1962. However, this parasite was not observed in either \u003cem\u003eM. surmuletus\u003c/em\u003e or \u003cem\u003eU. moluccensis\u003c/em\u003e specimens. All three \u003cem\u003eOrtholinea\u003c/em\u003e specimens (E-7, E-9, and E-13) exhibited an identical 18S rDNA genotype, clustering with \u003cem\u003eOrtholinea mullusi\u003c/em\u003e (MF539825; Gurkanlı et al., 2018) in both ML and BI phylogenetic trees (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). The 18S rDNA sequence similarity between \u003cem\u003eO. mullusi\u003c/em\u003e and the newly obtained genotype was 99.3%. Rangel et al. [\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e] reported that interspecific variation in 18S rDNA within the genus \u003cem\u003eOrtholinea\u003c/em\u003e ranges from 1.65% to 29.1%, suggesting that the isolates in this lineage are likely conspecific specimens of \u003cem\u003eO. mullusi\u003c/em\u003e. Additionally, the close similarity in myxospore morphology and the agreement of morphometric data further support this conclusion (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). The only difference observed between the original description of \u003cem\u003eO. mullusi\u003c/em\u003e and the newly obtained specimens was the shape of the polar capsules, which were pyriform in the original description and slightly ovoidal in the new specimens. We believe this difference may be due to adaptation to different geographical locations, which show significant variation in certain physical parameters, such as sea temperature and salinity (renging from 15\u0026permil; to 20\u0026permil; in the Black Sea and approximatesly 39\u0026permil; in the Mediterranean) [\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e, \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e74\u003c/span\u003e]. Therefore, based on morphological, morphometric, and molecular phylogenetic evidence, we assign the newly obtained \u003cem\u003eOrtholinea\u003c/em\u003e specimens in this study to \u003cem\u003eO. mullusi\u003c/em\u003e G\u0026uuml;rkanlı, Okkay, \u0026Ccedil;ift\u0026ccedil;i, Yurakhno \u0026amp; \u0026Ouml;zer, 2018.\u003c/p\u003e \u003cp\u003eThe genus \u003cem\u003eOrtholinea\u003c/em\u003e Shulman, 1962 currently includes 27 valid species, most of which infect marine fish hosts [\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e]. Only a few exceptions, such as \u003cem\u003eO. fluviatilis\u003c/em\u003e, \u003cem\u003eO. africanus\u003c/em\u003e, \u003cem\u003eO. lauquen\u003c/em\u003e, and \u003cem\u003eO. sphaerocapsularae\u003c/em\u003e, have been reported from non-marine environments [\u003cspan additionalcitationids=\"CR76\" citationid=\"CR75\" class=\"CitationRef\"\u003e75\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e77\u003c/span\u003e]. Although the genus shows relatively low species diversity compared to other myxozoan genera, its members have a broad global distribution [\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e]. In T\u0026uuml;rkiye, five \u003cem\u003eOrtholinea\u003c/em\u003e species have been reported to date: \u003cem\u003eO. divergens\u003c/em\u003e (ex \u003cem\u003eParablennius sanguinolentus\u003c/em\u003e), \u003cem\u003eO. gobiusi\u003c/em\u003e (ex \u003cem\u003eNeogobius melanostomus\u003c/em\u003e), \u003cem\u003eO. orientalis\u003c/em\u003e (ex \u003cem\u003eAlosa tanaica\u003c/em\u003e and \u003cem\u003eMullus barbatus\u003c/em\u003e), \u003cem\u003eO. mullusi\u003c/em\u003e (ex \u003cem\u003eMullus barbatus\u003c/em\u003e), and \u003cem\u003eO. hamsiensis\u003c/em\u003e (ex \u003cem\u003eEngraulis encrasicolus\u003c/em\u003e) [\u003cspan additionalcitationids=\"CR23\" citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e]. All of these species have been recorded from the Black Sea coasts of T\u0026uuml;rkiye. However, to date, no \u003cem\u003eOrtholinea\u003c/em\u003e species have been reported from the Sea of Marmara or along the Turkish coasts of the Aegean and Mediterranean Seas. \u003cem\u003eOrtholinea mullusi\u003c/em\u003e, the species identified in this study, was originally described from \u003cem\u003eM. barbatus\u003c/em\u003e along the Black Sea coasts of T\u0026uuml;rkiye and has not been reported elsewhere [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. Therefore, this study represents the second global record for this species. Given its occurrence in the Black Sea and the Mediterranean Sea, it is plausible that \u003cem\u003eO. mullusi\u003c/em\u003e also infects \u003cem\u003eM. barbatus\u003c/em\u003e populations in the Aegean and Marmara Seas. However, further research is needed to confirm this. Species of the genus \u003cem\u003eOrtholinea\u003c/em\u003e are mainly coelozoic and have predominantly been reported from organs of the urinary system [\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e]. Consistent with these data, \u003cem\u003eO. mullusi\u003c/em\u003e was originally described from the urinary bladder and kidney [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. Similarly, in the present study, \u003cem\u003eO. mullusi\u003c/em\u003e specimens E-9 and E-13 were observed in the kidney and urinary bladder, respectively. Additionally, specimen E-7 was detected in the gonad and the attached oviduct. Considering the coelozoic nature of the genus \u003cem\u003eOrtholinea\u003c/em\u003e, it is more likely that this specimen originated from the oviduct rather than the gonadal tissue. Likewise, another species of the genus, \u003cem\u003eO. undulans\u003c/em\u003e, has been reported from both the urinary system and the oviduct [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e]. G\u0026uuml;rkanlı et al. [\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e] reported an infection prevalence of \u003cem\u003eO. mullusi\u003c/em\u003e in \u003cem\u003eM. barbatus\u003c/em\u003e along the Black Sea coast of 24.5% (49 infected hosts out of 200 examined fish). In contrast, in this study, this parasite was detected in only 3 of the 330 examined \u003cem\u003eM. barbatus\u003c/em\u003e individuals (0.9%). Moreover, as mentioned earlier, it was not observed in any specimens of \u003cem\u003eM. surmuletus\u003c/em\u003e or \u003cem\u003eU. moluccensis\u003c/em\u003e. Additionally, \u003cem\u003eO. mullusi\u003c/em\u003e was detected in \u003cem\u003eM. barbatus\u003c/em\u003e specimens collected from Muğla (the western Mediterranean coast of Turkiye) and Antalya (the central Mediterranean coast of Turkiye), but no infection was recorded in samples from Mersin (the eastern Mediterranean coast of Turkiye). The significant difference in infection rates between the Black Sea and Mediterranean populations, along with the limited distribution of the parasite in the Mediterranean Sea, suggests that \u003cem\u003eO. mullusi\u003c/em\u003e may have only recently been introduced into the Mediterranean Sea. The species\u0026rsquo; adaptation and expansion processes could still be ongoing.\u003c/p\u003e \u003cp\u003eIn this study, three mullid hosts were examined for myxozoan infections; however, no myxozoan parasites were observed in \u003cem\u003eU. moluccensis\u003c/em\u003e. This host is a Lessepsian species, with the earliest records along the eastern Mediterranean coast dating back to the 1947 [\u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e]. Although \u003cem\u003eU. moluccensis\u003c/em\u003e has a wide native distribution, extending from the Red Sea, East Africa, Madagascar and R\u0026eacute;union east to the Caroline Islands and New Guinea, north to southern Japan, and southward to Western Australia and Queensland (Australia), there are no documented records of myxozoan infections for this species [\u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e81\u003c/span\u003e]. However, one myxozoan parasite, \u003cem\u003eCeratomyxa sultani\u003c/em\u003e, has been reported from another \u003cem\u003eUpeneus\u003c/em\u003e species, \u003cem\u003eU. margarethae\u003c/em\u003e, in the Arabian Gulf, suggesting that \u003cem\u003eU. moluccensis\u003c/em\u003e may also harbor myxosporean parasites in its native range [\u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e]. Upon migration to the eastern Mediterranean, this fish may have lost its myxosporean parasites, potentially due to the absence of the appropriate primary invertebrate host. Future studies incorporating various ecological factors are needed to further elucidate this phenomenon.\u003c/p\u003e \u003cp\u003eThe main outcomes of this study are summarised as follows: \u003cb\u003ea)\u003c/b\u003e a new myxosporean species, named \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp., was identified from the trunk muscles of \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e; \u003cb\u003eb)\u003c/b\u003e this species is the second \u003cem\u003eKudoa\u003c/em\u003e species known to infect the Mullidae family worldwide and the first \u003cem\u003eKudoa\u003c/em\u003e species recorded from Mullidae in the Mediterranean Sea; \u003cb\u003ec)\u003c/b\u003e a second new myxosporean species, named \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp., was identified in the gall bladder of \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e; \u003cb\u003ed)\u003c/b\u003e this species is the second \u003cem\u003eMyxobolus\u003c/em\u003e species, after \u003cem\u003eM. parvus\u003c/em\u003e, reported to infect \u003cem\u003eM. barbatus\u003c/em\u003e globally; \u003cb\u003ee)\u003c/b\u003e this study provides the first record of \u003cem\u003eO. mullusi\u003c/em\u003e from the Mediterranean Sea; \u003cb\u003ef)\u003c/b\u003e unexpectedly, no myxosporean infections were found in \u003cem\u003eU. moluccensis\u003c/em\u003e.\u003c/p\u003e"},{"header":"Declarations","content":" \u003ch2\u003eCompeting interests\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no conflict of interest.\u003c/p\u003e \u003ch2\u003eEthical standards\u003c/h2\u003e \u003cp\u003eAll applicable international, national, and institutional guidelines for the care and use of animals were followed.\u003c/p\u003e \u003ch2\u003eFinancial support\u003c/h2\u003e \u003cp\u003eThis study received no grant from any funding agency.\u003c/p\u003e \u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eE. \u0026Ouml;zdemir provided support for the fieldwork of the study and initial laboratory studies, including dissections and parasitological examinations. For molecular work, C.T. G\u0026uuml;rkanlı and E. \u0026Ouml;zdemir obtained DNA sequence data from parasites and conducted the phylogenetic analyses. C.T. G\u0026uuml;rkanlı and E. \u0026Ouml;zdemir prepared the firs draft of the manuscript. C.T. G\u0026uuml;rkanlı oversaw all subsequent revisions of the manuscript and wrote the final text. All authors reviewed the manuscript and provided edits on the final version.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e \u003cp\u003eThe authors gratefully acknowledge Prof. Dr. Ahmet \u0026Ouml;zer for his valuable contributions to the finalization of the manuscript.\u003c/p\u003e\u003ch2\u003eData availability statement\u003c/h2\u003e \u003cp\u003eSequence data is available on the NCBI GenBank database. 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Saudi J Biol Sci 25:597\u0026ndash;603\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"acta-parasitologica","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"actp","sideBox":"Learn more about [Acta Parasitologica](http://link.springer.com/journal/11686)","snPcode":"11686","submissionUrl":"https://submission.springernature.com/new-submission/11686/3","title":"Acta Parasitologica","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Goatfish, Kudoa, Ortholinea, Myxobolus, Mediterranean, Türkiye","lastPublishedDoi":"10.21203/rs.3.rs-9538756/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9538756/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eMyxozoa are globally distributed microscopic parasites that primarily infect fish but also occur in amphibians and, more rarely, reptiles, birds, mammals, and annelid worms. The aim of this study was to determine the diversity and parasitological characteristics of Myxozoa infecting \u003cem\u003eMullus barbatus\u003c/em\u003e, \u003cem\u003eMullus surmuletus\u003c/em\u003e, and \u003cem\u003eUpeneus moluccensis\u003c/em\u003e along the Mediterranean coasts of T\u0026uuml;rkiye. For this purpose, 330 \u003cem\u003eM. barbatus\u003c/em\u003e, 253 \u003cem\u003eM. surmuletus\u003c/em\u003e, and 240 \u003cem\u003eU. moluccensis\u003c/em\u003e specimens were collected from three localities representing the western (Muğla), central (Antalya), and eastern (Mersin) regions of the Turkish Mediterranean coast and were subjected to parasitological investigations. Based on morphological and molecular analyses, three myxozoan species-\u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp., \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp., and \u003cem\u003eOrtholinea mullusi\u003c/em\u003e-were identified, two of which represent novel species. \u003cem\u003eKudoa mediterraneus\u003c/em\u003e n. sp., infecting the trunk muscles mainly of \u003cem\u003eM. barbatus\u003c/em\u003e, was identified as a new species. This species represents the second \u003cem\u003eKudoa\u003c/em\u003e species reported from the family Mullidae worldwide and the first record of a \u003cem\u003eKudoa\u003c/em\u003e species infecting Mullidae in the Mediterranean Sea. The second novel species identified in this study, \u003cem\u003eMyxobolus mullus\u003c/em\u003e n. sp., infects the gall bladder of \u003cem\u003eM. barbatus\u003c/em\u003e and \u003cem\u003eM. surmuletus\u003c/em\u003e. This species represents the second \u003cem\u003eMyxobolus\u003c/em\u003e species, after \u003cem\u003eMyxobolus parvus\u003c/em\u003e, reported to infect \u003cem\u003eM. barbatus\u003c/em\u003e worldwide. \u003cem\u003eOrtholinea mullusi\u003c/em\u003e, originally described from \u003cem\u003eM. barbatus\u003c/em\u003e in the Black Sea, was recorded as the third myxozoan species in the present study. It was observed in the gonad, urinary bladder, and kidney of three \u003cem\u003eM. barbatus\u003c/em\u003e specimens. Unexpectedly, no myxosporean infections were detected in \u003cem\u003eU. moluccensis\u003c/em\u003e.\u003c/p\u003e","manuscriptTitle":"Diversity and Parasitological Traits of Myxozoan Parasites Infecting Various Mullidae Species Along the Mediterranean Coasts of Türkiye","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-05-13 13:43:14","doi":"10.21203/rs.3.rs-9538756/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"198640891636701334981261282212223372117","date":"2026-05-06T09:51:21+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-05-05T12:26:40+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-27T16:18:28+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-27T16:18:08+00:00","index":"","fulltext":""},{"type":"submitted","content":"Acta Parasitologica","date":"2026-04-27T08:28:56+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"acta-parasitologica","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"actp","sideBox":"Learn more about [Acta Parasitologica](http://link.springer.com/journal/11686)","snPcode":"11686","submissionUrl":"https://submission.springernature.com/new-submission/11686/3","title":"Acta Parasitologica","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"b67d2db8-7947-4ef9-a981-1c934f118350","owner":[],"postedDate":"May 13th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewerAgreed","content":"198640891636701334981261282212223372117","date":"2026-05-06T09:51:21+00:00","index":20,"fulltext":""},{"type":"reviewersInvited","content":"9","date":"2026-05-05T12:26:40+00:00","index":"","fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-13T13:43:14+00:00","versionOfRecord":[],"versionCreatedAt":"2026-05-13 13:43:14","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9538756","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9538756","identity":"rs-9538756","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}