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A synopsis of 43 nominal species from five genera of tailed Myxobolidae infecting Indian freshwater and marine fishes is presented. The main characteristic of this group is the presence of at least one tail-like caudal process. For each species, relevant morphological and morphometric data are provided, such as the host(s), site(s) of infection within the host and sampling state. A key for the identification of 13 genera of tailed Myxobolidae is also included.

Key to tailed genera of Myxobolidae 1a. Spores ovoid or pyriform, with two polar capsules and a single tail-like caudal process................................. 2 1b. Spores with two or more tail-like caudal processes or bifurcated tail; composed of same materials as shell valve......3 2a. Single tail-like caudal process; caudal process symmetrically located at the posterior end........................................................................................ Unicauda Davis, 1944 2b. Single tail-like caudal process, which may be forked at the very end, originating posterolaterally and asymmetrically.................. Laterocaudata Chen et Hsieh, 1984 3a. Spores tear-shaped shaped, with a single polar capsule, caudal process bifurcated..... Phlogospora Qadri, 1962 3b. Spores with two polar capsules and 2 caudal processes... 4 3c. Spores with two polar capsules and 4 caudal processes... 5 4a. Spores with two caudal processes.......................................................................... Henneguya Thélohan, 1892 4b. Spores with two caudal processes, asymmetric...................... Hennegoides Lom, Tonguthai et Dyková, 1991 4c. Spores with two caudal processes, extending in opposite directions............... Dicauda Hoffman et Walker, 1978 5a. Spores ellipsoidal with two caudal processes at each end of the spore; polar capsules situated asymmetrically............................................. Neohenneguya Tripathi, 1953 5b. Spores with four posterolateral caudal processes, two from each shell valve........................................................................... Tetrauronema Wu, Wang et Jiang, 1988 5c. Spores rhomboidal with four caudal processes pointing in the opposite direction, connected by transverse filaments............................ Trigonosporus Hoshina, 1952

Myxobolus opsaridiumi sp. nov. urn:lsid:zoobank.org:act: F3105EFD-A1BC-483F-8FC8-5D07FC9B82F0 Figs 1���4; Table 1 Etymology The specific epithet is related to the host genus name. Type material CAMEROON ��� infected skin, muscle and spleen of Opsaridium ubangiensis with plasmodia; Centre Region, Anga River, Yaounde; deposited in parasitological collection of the Zoology Department Museum, College of Science, King Saud University, Saudi Arabia; Myxospsar/12/2018. Taxonomic summary Type host Opsaridium ubangiense Pellegrin, 1901 (Cyprinidae). Infected tissues Skin, muscles and spleen. Prevalence 54.7% (288 parasitized fish out of 526 examined). Vegetative stages Ovoid, spherical or ellipsoid plasmodia, variable in size, measuring from 0.3 mm to 2.5 mm in length and 0.2 mm to 1.5 mm in width. Description of myxospores (Fig. 1) Mature spores were ovoid to subspherical in frontal view and lenticular in lateral view (Fig. 1 A���B). The valves were relatively thick, without edge markings. Intercapsular processes were absent. The spore size was 10.7 �� 0.14 (10���11.5) ��m long, 9 ��0.15 (8���10) ��m wide and 6.2�� 0.7 (5.6���7.2) ��m thick. The two ovoid polar capsules were equal in size, converging and opening together at the anterior end of the same pore (Fig. 1A, C���D). They measured 5�� 0.07 (4.3���6.0) ��m in length and 2.7�� 0.07 (2.2���3.0) ��m in width. Polar filaments were coiled from 5 to 7 turns perpendicular to the longitudinal axis of the polar capsules (Fig. 1D). A sporoplasm containing an iodinophilous vacuole of varying shape and size filled the entire space below the polar capsules (Fig. 1A). Clinical finding and histopathology Based solely on gross observation of the fish, no signs of disease were observed. Parasitized fish harbored cysts on skin, muscles and spleen. On skin, white cysts up to 2 mm long were collected from the body flanks of some fish (Fig. 2A). Sections revealed that plasmodia developed in the connective tissue of the dermis beneath the underside of scales (Fig. 2B). Plasmodia were flattened and surrounded by a thin membrane and an internal endoplasm comprising a loosely defined matrix containing developed spores (Fig. 2B). Some plasmodia were spotted within muscle cells (Fig. 2C). Plasmodia were spindle-shaped, centrally located in the cell and not surrounded by a visible wall. No evidence of inflammation or immune-cell recruitment was seen. The integrity of myofibrils within the infected fibers showed some degree of lysis, with partial loss of myofibrillar details and striations (Fig. 2 C���D). These lesions were observed close to plasmodia. Mature spores were scattered in the cytoplasm of infected cells (Fig. 2D). Infected spleens had plasmodia of up to 2.5 �� 1.5 mm (Fig. 3A). They were white, isolated or clustered (Fig. 3B). Some infected spleens were heavily infected and plasmodia were randomly distributed in the whole organ. In these cases, abnormal enlargement of the spleen was evident (Fig. 3C). Histological sections revealed that, for moderately infected spleens, cysts were either fixed to the external region of the organ (Fig. 3D) or completely implanted within it (Fig. 3E). Development of cysts in the spleen was asynchronous (Fig. 3 D���F). Atrophy of the adjacent splenic cells surrounding the cyst was likely due to mechanical compression (Fig. 3 F���G). Each plasmodium was surrounded by a wall of a monolayer of flat cells (Fig. 3 F���G). The central part of the plasmodium was occupied by fully mature spores, with initial stages of development visible in the periphery (Fig. 3G). * Refers to the largest polar capsule. ** Refers to the smallest polar capsule. Phylogenetic position Partial SSU rDNA sequences obtained fromdifferent organs were 100% identical.The consensus sequence of 1667 base pairs was submitted to GenBank with the accession number MN 497413. This sequence did not match any publicly available myxozoan sequence. The sequence with the highest nucleotide similarity, at 91.8%, was for Myxobolus haichengensis Chen, 1958 (GenBank entry KY965936), which reportedly infects the gills of Abbottina rivularis (Basilewsky, 1855). Similarity with M. dibombensis Folefack et al., 2019, a species we recently sequenced from Labeobarbus batesii (Boulenger, 1903) in Cameroon, was only 88.6%. The phylogenetic position of the newly sequenced species was analyzed with maximum likelihood and Bayesian inference methods. Both methods produced an identical topology. Myxobolus opsaridiumi sp. nov. occurs in a large clade that includes species infecting cyprinids (Fig. 4). The new species exhibits the highest phylogenetic affinity with M. haichengensis, Myxobolus sp. (accession number MG 253819) from the fins of Capoeta tinca (Heckel, 1843) off Anatolia, and M. squamae Keysselitz, 1908 infecting the skin of the common barbel Barbus barbus (Linnaeus, 1758)., Published as part of Lekeufack-Folefack, Guy Benoit, Tchoutezo-Tiwa, Armandine Estelle, Fomena, Abraham & Mansour, Lamjed, 2021, Myxobolus opsaridiumi sp. nov. (Cnidaria: Myxosporea) infecting different tissues of an ornamental fish, Opsaridium ubangiensis (Pellegrin, 1901), in Cameroon: morphological and molecular characterization, pp. 56-71 in European Journal of Taxonomy 733 on pages 59-64, DOI: 10.5852/ejt.2021.733.1221, http://zenodo.org/record/5706193, {"references":["Basu S. & Haldar D. 2003. Three new species of Myxobolus Butschli, 1882 from different food fishes of West Bengal, India. Acta Protozoologica 42: 245 - 251.","Ali M. A., Al-Rasheid K. A., Sakran T., Abdel-Baki A. - A. & Abdel-Ghaffar F. A. 2002. Some species of the genus Myxobolus (Myxozoa: Myxosporea) infecting freshwater fish of the River Nile, Egypt, and the impact on their hosts. Parasitology Research 88: 9 - 15. https: // doi. org / 10.1007 / s 004360100449","Mandour A., Galal A. & Abed G. 1993. Myxobolus clarii n. sp. in the testis of the fish Clarias lazera from the River Nile of Assiut [Egypt]. Assiut Veterinary Medical Journal 29 (58): 108 - 114.","Eiras J. C., Monteiro C. M & Brasil-Sato M. C. 2010. Myxobolus franciscoi sp. nov. (Myxozoa: Myxosporea: Myxobolidae), a parasite of Prochilodus argenteus (Actinopterygii: Prochilodontidae) from the Upper Sao Francisco River, Brazil, with a revision of Myxobolus spp. from South America. Zoologia (Curitiba) 27 (1): 131 - 137.","Reed C. C., Basson L. & Van As L. L. 2003. Myxozoans infecting the sharptooth catfish, Clarias gariepinus in the Okavango River and Delta, Botswana, including descriptions of two new species, Henneguya samochimensis sp. n. and Myxobolus gariepinus sp. n. Folia Parasitologica (Praha) 50 (3): 183 - 189. https: // doi. org / 10.14411 / fp. 2003.033","Boungou M., Kabre G. B., Sakiti N. G., Marques A. & Sawadogo L. 2006. Description of four new myxosporean species (Myxozoa: Myxosporea) from genus Myxobolus, fish parasites of Burkina Faso, West Africa. Journal of Biological Sciences 6 (5): 861 - 867.","Eiras J. C., Molnar K. & Lu Y. S. 2005. Synopsis of the species of Myxobolus Butschli, 1882 (Myxozoa: Myxosporea: Myxobolidae). Systematic Parasitology 61 (1): 1 - 46. https: // doi. org / 10.1007 / s 11230 - 004 - 6343 - 9","Sakiti G. N. 1997. Myxosporidia and Microsporidia Fish Parasites in Southern Benin: Faunal Description, Ultrastructure and Biology. PhD thesis. National University of Benin.","Haldar D. P., Das M. K. & Sharma B. K. 1983. Studies on protozoan parasites from fishes: four new species of the genera Henneguya Thelohan, 1892, Thelohanellus Kudo, 1933, and Myxobolus Butschli, 1892. Archiv fur Protistenkunde 127 (3): 283 - 296. https: // doi. org / 10.1016 / S 0003 - 9365 (83) 80023 - 2","Abakar-Ousman A., Fomena P., Ngassam & Bouix G. 2006. Myxosporidian (Myxozoa) parasites of fresh water fish teleostean from Chad: new or few known species. Annales de l'Universite de N'Djamena, Ser. C 1: 111 - 121."]}

Lekeufack-Folefack, Guy Benoit, Tchoutezo-Tiwa, Armandine Estelle, Fomena, Abraham, Mansour, Lamjed (2021): Myxobolus opsaridiumi sp. nov. (Cnidaria: Myxosporea) infecting different tissues of an ornamental fish, Opsaridium ubangiensis (Pellegrin, 1901), in Cameroon: morphological and molecular characterization. European Journal of Taxonomy 733: 56-71, DOI: https://doi.org/10.5852/ejt.2021.733.1221

Myxobolus jialingensis n. sp. http://zoobank.org/ urn:lsid:zoobank.org:pub: 6410134B-D7AF-4C5C-920E-CC16A02BD7C9 ( Fig. 1 ), Published as part of Gao, Lei, Zhang, Jing, Yang, Chengzhong & Zhao, Yuanjun, 2020, Myxobolus jialingensis n. sp. (Myxozoa: Myxobolidae) infecting urinary bladder and hepatopancreas of yellowhead catfish Tachysurus fulvidraco from China, pp. 179-186 in Zootaxa 4819 (1) on page 180, DOI: 10.11646/zootaxa.4819.1.10, http://zenodo.org/record/3955802

Genus Myxobolus B��tschli, 1882 Taxonomic summary Type host: Tachysurus fulvidraco (Richardson, 1846) (Actinopterygii: Siluriformes: Bagridae). Type locality: Jialing River (29��58' N, 106��45' E), Shapingba District, Chongqing, China. Sites of infection: urinary bladder, hepatopancreas. Date of sampling: July 10, 2014. Prevalence: Of the 70 fish examined, two were infected (2.9%). Deposition of type materials: A syntype (mounted in glycerin-alcohol-formalin) has been deposited in the Collection Center of Chongqing Key Laboratory of Animal Biology, Chongqing Normal University, Chongqing, China (Coll. No. cq-2014071001). The DNA sample has been deposited in the same location, with the code number of cq-2014071002. Etymology: The species was named from the river (Jialing River), where the host was collected. Morphological description. A single white oval cyst (3.1 �� 2.8 mm) was discovered from hepatopancreas of a fish, and a white sphere cyst with a diameter of 3 mm detected in urinary bladder of another fish. Vegetative stages were not encountered. No obvious pathological traits for the host were observed. Therefore, the pathological damage to the fish remained unknown. The myxospores that infected the urinary bladder had a membranous sheath, and the spores from hepatopancreas had no (Fig. 1). The morphology of the spore body (excluding the membranous sheath) from different infection sites were the same. The mature spore was pyriform with a slightly pointed anterior and blunt posterior, which was 15.8 �� 0.7 (15.4���17.0) ��m in length and 8.0 �� 0.3 (7.8���8.9) ��m in width (n=37). Two pyriform polar capsules were slightly unequal in size, with the larger one of 7.4 �� 0.3 (6.7���8.0) ��m in length and 3.1 �� 0.2 (2.8���3.6) ��m in width (n=37), and the smaller one of 7.3 �� 0.3 (6.6���8.1) ��m in length and 3.3 �� 0.2 (2.9���3.6) ��m in width (n=37). Polar filaments coiled with 7���8 turns (Table 1). Remarks. Of the approximately 900 species of Myxobolus described (Eiras et al. 2005, 2014; Lom and Dykov�� 2006), M. jialingensis n. sp. has displayed similarity to Myxobolus voremkhai (Akhmerov, 1960), Myxobolus pseudowulii Zhang et al., 2017 and Myxobolus xiantaoensis Tahir et al., 2019, based on the fact that they all have pyriform spores with two unequal polar capsules (Table 1). Also, the four species all parasitize on/in the same host, Tachysurus fulvidraco. However, they could be distinguished from each other by spore dimension, size of polar capsule, turns of filament coils and infection sites. The infection sites of the new species are urinary bladder and hepatopancreas rather than gills for M. voremkhai, skin for M. pseudowulii and fins for M. xiantaoensis. The spore and Abbreviations: SL, spore length; SW, spore width; PCL, polar capsule length; PCW, polar capsule width; NC, number of coils in polar filame. polar capsules of M. jialingensis n. sp. are slightly larger than that of M. voremkhai (15.4���17.0 ��m �� 7.8���8.9 ��m vs. 13.1���15.5 ��m �� 7.1���8.6 in length and width of spores; 6.7���8.0 ��m �� 2.8���3.6 ��m vs. 6.3���7.4 ��m �� 2.2���3.3 in length and width of the larger polar capsules; 6.6���8.1 ��m �� 2.9���3.6 ��m vs. 5.8���7.1 ��m �� 2.2���3.3 in length and width of the smaller polar capsules) (Table 1). And the number of polar filament coils for M. jialingensis n. sp. is much more than that of M. voremkhai (7���8 vs. 5���7). The new species has longer spores than M. pseudowulii (15.4���17.0 ��m vs. 12.9���16.2 ��m), and its proportion of the polar capsule in the whole spore is smaller than that of M. pseudowuli. The spore of M. jialingensis n. sp. is obviously longer and slightly narrower than that of M. xiantaoensis (15.4���17.0 ��m vs. 11.3���13.9 ��m in length; 7.8���8.9 ��m vs. 8.4���11.1 ��m in width). The new species is somewhat similar to M. tunicatusb and M. macrocapsularis. They all infect urinary blad- der of T. fulvidraco. However, the new species could be distinguished from the later two by its two unequal polar capsules (vs. two equal size for M. tunicatusb and M. macrocapsularis). And the morphometrics of the three species are apparently different (Table 1)., Published as part of Gao, Lei, Zhang, Jing, Yang, Chengzhong & Zhao, Yuanjun, 2020, Myxobolus jialingensis n. sp. (Myxozoa: Myxobolidae) infecting urinary bladder and hepatopancreas of yellowhead catfish Tachysurus fulvidraco from China, pp. 179-186 in Zootaxa 4819 (1) on pages 181-183, DOI: 10.11646/zootaxa.4819.1.10, http://zenodo.org/record/3955802, {"references":["Eiras, J., Molnar, K. & Lu, Y. S. (2005) Synopsis of the species of Myxobolus Butschli, 1882 (Myxozoa: Myxosporea: Myxobolidae). Systematic Parasitology, 61 (1), 1 - 46. https: // doi. org / 10.1007 / s 11230 - 004 - 6343 - 9","Eiras, J., Zhang, J. Y. & Molnar, K. (2014) Synopsis of the species of Myxobolus Butschli, 1882 (Myxozoa: Myxosporea, Myxobolidae) described between 2005 and 2013. Systematic Parasitology, 88 (1), 11 - 36. https: // doi. org / 10.1007 / s 11230 - 014 - 9484 - 5","Lom, J. & Dykova, I. (2006) Myxozoan genera: definition and notes on taxonomy, life-cycle terminology and pathogenic species. Folia Parasitologica, 53 (1), 1 - 36. https: // doi. org / 10.14411 / fp. 2006.001","Zhang, B., Zhai, Y. H., Liu, Y. & Gu, Z. M. (2017) Myxobolus pseudowulii sp. n. (Myxozoa: Myxosporea), a new skin parasite of yellow catfish Tachysurus fulvidraco (richardson) and redescription of Myxobolus voremkhai (Akhmerov, 1960). Folia Parasitologica, 64, 30. https: // doi. org / 10.14411 / fp. 2017.030"]}

Myxobolus jialingensis n. sp. http://zoobank.org/ urn:lsid:zoobank.org:pub: 6410134B-D7AF-4C5C-920E-CC16A02BD7C9 ( Fig. 1 )

Myxobolus adrianoi sp. nov . urn:lsid:zoobank.org:act: 9FF87134-46B5-47AD-98FE-B97 E 6F913645 Figs 1���4 Etymology The specific name, adrianoi, is in homage to Dr. Edson Adriano, Professor at Federal University of S��o Paulo, Brazil, who has been contributing to improving our knowledge on the diversity of South American Myxosporea. Material examined Syntypes BRAZIL ��� 10+ myxospores; Amazonas State, L��brea Municipality; 7��15���32��� S, 64��47���52��� W; air-dried slide stained with Giemsa; MZUSP 8469. Type host Corydoras schwartzi R��ssel, 1963 (Siluriformes: Callichthyidae). Site of infection Serosa layer of intestine. Prevalence From 30 examined fish, five were infected (16.6%). Description Mature myxospores ovoid in body shape in frontal view, showing two aubergine-shaped, elongate symmetrical polar capsules occupying more than half length of spore. Total myxospore length 22.4 �� 0.3 ��m and width 16.3 �� 0.1 ��m. Two polar capsules, 14.3 �� 0.2 ��m in length and 6.5 ��0.1 ��m in width (Figs 2 A���D, 3). Ultrastructural analysis showed myxospores in various stages of development. In young developmental myxospore stage, the polar filament could still be observed out of the polar capsules and valvogenic cells were readily recognized by valve-forming materials (Fig. 4A). In myxospores at a more advanced developmental stage, a polar filament inside the polar capsule and binucleated sporoplasms containing a moderate number of sporoplasmosomes were observed (Fig. 4 B���C). Transverse sections of a polar capsule provided evidence of a polar filament with five coils (Fig. 4 B���C). Sections of immature myxospores show the valvogenic cells abutting each other to form a sutural ridge, and frequently the two cells were separated by a thin layer of homogenous material (Fig. 4D). Almost mature myxospores showed sutural lines, valve-forming material, valves, two nuclei and sporoplasmosomes at sporoplasms (Fig. 4 E���F). The outer surface of the myxospore valves was smooth, with no evidence of formation of ridges in the valves (Fig. 4 E���F)., Published as part of Mathews, Patrick D., Madrid, Rafael R. M., Mertins, Omar, Rigoni, Vera L. S. & Morandini, Andr�� C., 2020, A new Myxobolus (Cnidaria: Myxosporea) infecting the ornamental catfish Corydoras schwartzi from the Purus River in Brazil, pp. 1-14 in European Journal of Taxonomy 620 on pages 4-6, DOI: 10.5852/ejt.2020.620, http://zenodo.org/record/3750231

Myxobolus adrianoi sp. nov . urn:lsid:zoobank.org:act: 9FF87134-46B5-47AD-98FE-B97 E 6F913645 Figs 1–4 Etymology The specific name, adrianoi, is in homage to Dr. Edson Adriano, Professor at Federal University of São Paulo, Brazil, who has been contributing to improving our knowledge on the diversity of South American Myxosporea. Material examined Syntypes BRAZIL • 10+ myxospores; Amazonas State, Lábrea Municipality; 7°15′32″ S, 64°47′52″ W; air-dried slide stained with Giemsa; MZUSP 8469. Type host Corydoras schwartzi Rössel, 1963 (Siluriformes: Callichthyidae). Site of infection Serosa layer of intestine. Prevalence From 30 examined fish, five were infected (16.6%). Description Mature myxospores ovoid in body shape in frontal view, showing two aubergine-shaped, elongate symmetrical polar capsules occupying more than half length of spore. Total myxospore length 22.4 ± 0.3 μm and width 16.3 ± 0.1 μm. Two polar capsules, 14.3 ± 0.2 μm in length and 6.5 ±0.1 μm in width (Figs 2 A–D, 3). Ultrastructural analysis showed myxospores in various stages of development. In young developmental myxospore stage, the polar filament could still be observed out of the polar capsules and valvogenic cells were readily recognized by valve-forming materials (Fig. 4A). In myxospores at a more advanced developmental stage, a polar filament inside the polar capsule and binucleated sporoplasms containing a moderate number of sporoplasmosomes were observed (Fig. 4 B–C). Transverse sections of a polar capsule provided evidence of a polar filament with five coils (Fig. 4 B–C). Sections of immature myxospores show the valvogenic cells abutting each other to form a sutural ridge, and frequently the two cells were separated by a thin layer of homogenous material (Fig. 4D). Almost mature myxospores showed sutural lines, valve-forming material, valves, two nuclei and sporoplasmosomes at sporoplasms (Fig. 4 E–F). The outer surface of the myxospore valves was smooth, with no evidence of formation of ridges in the valves (Fig. 4 E–F).

Myxobolus arariensis n. sp. ( Figs 1���3) Morphological description. Mature spores are ellipsoidal in shape, with a mean length of 11.4 ��m (10.7��� 12.6) and mean width of 7.2 ��m (6.4���7.9). Each spore contains two polar capsules (PCs) of equal size 4.0 �� 0.7 ��m (3.6���4.3) long and 1.9 �� 0.36 ��m (1.7���2.2) in width (Figs. 1B and 3) (Table 1). Type host. Rhamdia quelen (Quoy & Gaimard 1824) Site of infection. epaxial and hypaxial layers of the musculature, with plasmodia containing numerous spores. Type-locality. Arari River, Cachoeira do Arari on Maraj�� Island, northern Brazil. Etymology. The species was named for the locality of origin, the Arari River in northern Brazil. Specimens deposited. Microscope slides containing spores from the muscle layer, prepared using the paraffin technique, stained in Gutierrez and mounted in Entellan were deposited in the International Protozoan Type Collection of the National Institute for Amazonian Research (INPA) in Manaus, Amazonas state, Brazil (catalog number: INPA /027). The partial 18S rDNA sequence was deposited in GenBank under accession number MG572219. Prevalence. Three of twenty-five R. quelen examined 12% (3/25) had plasmodia of an unknown parasite from the genus Myxobolus. Remarks. M. arariensis can be differentiated morphologically from all seven Myxobolus species known to infect the muscle tissue of freshwater fish (Table 1). The new species can be distinguished from M. tasikkenyirensis (Sz��kely et al. 2009a) and M. groenlandicus (Buchmann et al. 2012) by the different shape of the anterior extremity of the spores, and from M. leptobarbi (Sz��kely et al. 2009b) by the same trait. The length of M. arariensis (11.4 ��m) is most similar to that of M. lentisuturalis (Dykov�� et al. 2002), which is 11.8 ��m long, whereas M. omari (Sz��kely et al. 2009b) is the shortest species, at 7.9 ��m. Anomalous spores with a caudal filament and lack of ornamentation on the external wall were also observed (Fig. 2C). Histology. The histological analysis revealed the presence of cysts of M. arariensis lodged in the fibers of the skeletal muscles (Fig. 4). Immature spores were observed in the most external layer of the cyst, with mature spores being found more internally. The cyst wall is thick and fibrous, and the adjacent musculature was compressed, with the sarcoplasm frayed, and evidence of a necrotic reaction caused by this compression. Molecular data. In the molecular analysis, the specific pair of myxozoan primers (MC5-MC3) amplified 974 bps of the 18S rDNA gene of the spores obtained from the plasmodia found infecting the musculature of R. quelen. The BLAST search of the 18S rDNA sequence data (974 bps) of the Myxobolus species parasitizing R. quelen found no identical myxozoan sequence in GenBank, although a similarity of at least 85% was found with four species: Myxobolus cordeiroi (KF296353, 90% similarity), Myxobolus sp. GA2 (KU 170935, 86%), Myxobolus lentisuturalis (AY278563, 85%), and Myxobolus cultus (HQ613409, 85%). The optimal evolutionary model for maximum likelihood (ML) and Bayesian analysis were determined by jModelTest 3.0 (Posada, 2008) which identified the best evolutionary model as the general time reversible model (GTR + I + G), using Akaike information criteria. Nucleotide frequencies were estimated from the data (A = 0.2574, C = 0.1848, G = 0.2625, T = 0.2326) and six rates of nucleotide substitution calculated as AC = 0.8659, AG = 2.6388, AT = 1.7658, CG = 0.4883, CT = 3.4814, GT = 1.000. The proportion of invariable site was 0.5565 and the alpha value of gamma distribution parameter 0.3612. Two independent runs were conducted with 4 chains for 2 million generations for Bayesian analysis. Ceratomyxa shasta (AF001579) e C. amazonensis (KX236169) was designated as outgroup. Phylogenetic trees were sampled every 100 generation. Characteristics of some Myxobolus species. Abbreviations: FC = Capsule Formate, SL, spore length, SW = Spore Width, PCL = Polar Capsule Length, PCW= Polar Capsule Width. PC = relative size of the polar capsules (= = equal in size, # = different in size, or equal and different); All measurements are given in micrometers. In the phylogenetic analysis, trees generated by Bayesian Inference (BI) had similar topologies, but with different support values at some nodes. A strong clustering tendency was found according to phylogenetic affinities. The phylogram indicated the existence of three clades, A, B and C the first paraphyletic, includes species of Henneguya and Myxobolus formed by the freshwater and marine water (Mugiliformes). The clade A subdivide into 2 subclades, A1 and A2. The subclade A1 shows M. arariensis grouping on the same branch with M. cordeiroi and with adjacent subclade with M. marajoensis species, having same host. The other subclade, A2 have the presence of the Myxobolus and Henneguya, corroborating the characteristic of the Myxobolus genus to be paraphyletic. On the other hand, the clades B and C have agrouped species of various Myxobolus parasites of freshwater fish belonging to same and different Order (Table 3). The type of host defines a well-supported freshwater and marine water clade of Myxobolus and Henneguya (clade A) that infect fish of the orders Siluriformes, Mugiliformes, Characiformes and Perciformes. However, M. arariensis, which infects silurids, evolved independently from the Myxobolus subclade A1 that infects fish of the families Pimelodidae and Ictaluridae. The tree presented a similar topology for the other clades, clustering according to the taxonomic order of the host. The clade B groups Myxobolus, of the orders Cypriniformes and Perciformes. The clade C is also compounded by Myxobolus groups that parasitize hosts of the different orders Salmoniformes, Siluriformes, Characiformes and Cypriniformes (Fig. 5). The p distance found between M. arariensis and any other Myxobolus species that had Siluriformes as host ranged from 11.6 to 22.1% (Table 2), which reinforces the definition of M. arariensis as a new species. The myxosporidian sequences analysed are showing in the Table 3. ������continued on the next page, Published as part of Abrunhosa, Jacqueline, Sindeaux-Neto, Jose L., Santos, Sidney, Hamoy, Igor & Matos, Edilson, 2018, A new species of myxozoa in the skeletal striated musculature of Rhamdia quelen (Quoy & Gaimard) (Siluriforme: Pimelodidae) Amazonian fish, Maraj�� island, Brazil, pp. 164-176 in Zootaxa 4482 (1) on pages 166-172, DOI: 10.11646/zootaxa.4482.1.8, http://zenodo.org/record/1453193, {"references":["Buchmann, K., Skovgaard, A. & Kania, P. W. (2012) Myxobolus groenlandicus n. sp. (Myxozoa) distorting skeletal structures and musculature of Greenland halibut Reinhardtius hippoglossoides (Teleostei: Pleuronectidae). Diseases of Aquatic Organisms, 98, 133 - 141. https: // dx. doi. org / 10.3354 / dao 02437"]}

Myxobolus bragantinus n. sp. Host: Mugil rubrioculus Harrison et al. 2007. Locality: Municipality of Bragan��a, northeastern Par��, Brazil. Site of infection: Gill filaments Plasmodial type: Intrafilamental-epithelial (FE) Prevalence: 60% (90/150). Type material: Slides containing cysts with spores in the gill filament, which were processed by embedding in paraffin, and stained with Ziehl-Neelsen (Fig. 2a). These specimens were deposited in the International Protozoan Type Specimen Collection at the Brazilian National Institute of Amazonian Research (INPA) in Manaus, Amazonas, Brazil (catalog number: INPA 34). Etymology: The species name, bragantinus, refers to the collection site, at Bragan��a, in the Brazilian state of Par��, where the microparasite was discovered and described for the first time. The histological analysis shows cysts lodged internally in the gill filament, located in the intermediate portion, causing hypertrophy, a moderate increase in the size of the filaments, associated with the presence of the parasitic cysts of Myxobolus bragantinus n. sp., and a thickening of the cyst wall (Fig. 2 a���b). Under light microscopy, the ZN-stained slides revealed spores characteristic of Myxobolus (Lom & Dykova 2006), featuring polar capsules (see Fig. 2b). The location of the parasite in the gills, is known as the ���intrafilamental-epithelial��� (FE) type., Published as part of Cardim, Joyce, Silva, Diehgo, Hamoy, Igor, Matos, Edilson & Abrunhosa, Fernando, 2018, Myxobolus bragantinus n. sp. (Cnidaria: Myxosporea) from the gill filaments of the redeye mullet, Mugil rubrioculus (Mugiliformes: Mugilidae), on the eastern Amazon coast, pp. 177-187 in Zootaxa 4482 (1) on pages 179-181, DOI: 10.11646/zootaxa.4482.1.9, http://zenodo.org/record/1440542, {"references":["Harrison, I. J., Nirchio, M., Oliveira, C., Ron, E. & Gaviria, J. (2007) A new species of mullet (Teleostei: Mugilidae) from Venezuela, with a discussion on the taxonomy of Mugil gaimardianus. Journal of Fish Biology, 71 (Supplement A), 76 - 97. https: // doi. org / 10.1111 / j. 1095 - 8649.2007.01520. x.","Lom, J. & Dikova, I. (2006) Myxozoan genera: definition and notes on taxonomy, life-cycle terminology and pathogenic species. Folia Parasitologica (Praha), 53 (1), 1 - 36. http: // dx. doi. org / 10.14411 / fp. 2006.001"]}

Myxobolus arariensis n. sp. ( Figs 1–3) Morphological description. Mature spores are ellipsoidal in shape, with a mean length of 11.4 µm (10.7– 12.6) and mean width of 7.2 µm (6.4–7.9). Each spore contains two polar capsules (PCs) of equal size 4.0 ± 0.7 µm (3.6–4.3) long and 1.9 ± 0.36 µm (1.7–2.2) in width (Figs. 1B and 3) (Table 1). Type host. Rhamdia quelen (Quoy & Gaimard 1824) Site of infection. epaxial and hypaxial layers of the musculature, with plasmodia containing numerous spores. Type-locality. Arari River, Cachoeira do Arari on Marajó Island, northern Brazil. Etymology. The species was named for the locality of origin, the Arari River in northern Brazil. Specimens deposited. Microscope slides containing spores from the muscle layer, prepared using the paraffin technique, stained in Gutierrez and mounted in Entellan were deposited in the International Protozoan Type Collection of the National Institute for Amazonian Research (INPA) in Manaus, Amazonas state, Brazil (catalog number: INPA /027). The partial 18S rDNA sequence was deposited in GenBank under accession number MG572219. Prevalence. Three of twenty-five R. quelen examined 12% (3/25) had plasmodia of an unknown parasite from the genus Myxobolus. Remarks. M. arariensis can be differentiated morphologically from all seven Myxobolus species known to infect the muscle tissue of freshwater fish (Table 1). The new species can be distinguished from M. tasikkenyirensis (Székely et al. 2009a) and M. groenlandicus (Buchmann et al. 2012) by the different shape of the anterior extremity of the spores, and from M. leptobarbi (Székely et al. 2009b) by the same trait. The length of M. arariensis (11.4 µm) is most similar to that of M. lentisuturalis (Dyková et al. 2002), which is 11.8 µm long, whereas M. omari (Székely et al. 2009b) is the shortest species, at 7.9 µm. Anomalous spores with a caudal filament and lack of ornamentation on the external wall were also observed (Fig. 2C). Histology. The histological analysis revealed the presence of cysts of M. arariensis lodged in the fibers of the skeletal muscles (Fig. 4). Immature spores were observed in the most external layer of the cyst, with mature spores being found more internally. The cyst wall is thick and fibrous, and the adjacent musculature was compressed, with the sarcoplasm frayed, and evidence of a necrotic reaction caused by this compression. Molecular data. In the molecular analysis, the specific pair of myxozoan primers (MC5-MC3) amplified 974 bps of the 18S rDNA gene of the spores obtained from the plasmodia found infecting the musculature of R. quelen. The BLAST search of the 18S rDNA sequence data (974 bps) of the Myxobolus species parasitizing R. quelen found no identical myxozoan sequence in GenBank, although a similarity of at least 85% was found with four species: Myxobolus cordeiroi (KF296353, 90% similarity), Myxobolus sp. GA2 (KU 170935, 86%), Myxobolus lentisuturalis (AY278563, 85%), and Myxobolus cultus (HQ613409, 85%). The optimal evolutionary model for maximum likelihood (ML) and Bayesian analysis were determined by jModelTest 3.0 (Posada, 2008) which identified the best evolutionary model as the general time reversible model (GTR + I + G), using Akaike information criteria. Nucleotide frequencies were estimated from the data (A = 0.2574, C = 0.1848, G = 0.2625, T = 0.2326) and six rates of nucleotide substitution calculated as AC = 0.8659, AG = 2.6388, AT = 1.7658, CG = 0.4883, CT = 3.4814, GT = 1.000. The proportion of invariable site was 0.5565 and the alpha value of gamma distribution parameter 0.3612. Two independent runs were conducted with 4 chains for 2 million generations for Bayesian analysis. Ceratomyxa shasta (AF001579) e C. amazonensis (KX236169) was designated as outgroup. Phylogenetic trees were sampled every 100 generation. Characteristics of some Myxobolus species. Abbreviations: FC = Capsule Formate, SL, spore length, SW = Spore Width, PCL = Polar Capsule Length, PCW= Polar Capsule Width. PC = relative size of the polar capsules (= = equal in size, # = different in size, or equal and different); All measurements are given in micrometers. In the phylogenetic analysis, trees generated by Bayesian Inference (BI) had similar topologies, but with different support values at some nodes. A strong clustering tendency was found according to phylogenetic affinities. The phylogram indicated the existence of three clades, A, B and C the first paraphyletic, includes species of Henneguya and Myxobolus formed by the freshwater and marine water (Mugiliformes). The clade A subdivide into 2 subclades, A1 and A2. The subclade A1 shows M. arariensis grouping on the same branch with M. cordeiroi and with adjacent subclade with M. marajoensis species, having same host. The other subclade, A2 have the presence of the Myxobolus and Henneguya, corroborating the characteristic of the Myxobolus genus to be paraphyletic. On the other hand, the clades B and C have agrouped species of various Myxobolus parasites of freshwater fish belonging to same and different Order (Table 3). The type of host defines a well-supported freshwater and marine water clade of Myxobolus and Henneguya (clade A) that infect fish of the orders Siluriformes, Mugiliformes, Characiformes and Perciformes. However, M. arariensis, which infects silurids, evolved independently from the Myxobolus subclade A1 that infects fish of the families Pimelodidae and Ictaluridae. The tree presented a similar topology for the other clades, clustering according to the taxonomic order of the host. The clade B groups Myxobolus, of the orders Cypriniformes and Perciformes. The clade C is also compounded by Myxobolus groups that parasitize hosts of the different orders Salmoniformes, Siluriformes, Characiformes and Cypriniformes (Fig. 5). The p distance found between M. arariensis and any other Myxobolus species that had Siluriformes as host ranged from 11.6 to 22.1% (Table 2), which reinforces the definition of M. arariensis as a new species. The myxosporidian sequences analysed are showing in the Table 3. ……continued on the next page

Abrunhosa, Jacqueline, Sindeaux-Neto, Jose L., Santos, Sidney, Hamoy, Igor, Matos, Edilson (2018): A new species of myxozoa in the skeletal striated musculature of Rhamdia quelen (Quoy & Gaimard) (Siluriforme: Pimelodidae) Amazonian fish, Marajó island, Brazil. Zootaxa 4482 (1): 164-176, DOI: https://doi.org/10.11646/zootaxa.4482.1.8

Myxobolus bragantinus n. sp. Host: Mugil rubrioculus Harrison et al. 2007. Locality: Municipality of Bragança, northeastern Pará, Brazil. Site of infection: Gill filaments Plasmodial type: Intrafilamental-epithelial (FE) Prevalence: 60% (90/150). Type material: Slides containing cysts with spores in the gill filament, which were processed by embedding in paraffin, and stained with Ziehl-Neelsen (Fig. 2a). These specimens were deposited in the International Protozoan Type Specimen Collection at the Brazilian National Institute of Amazonian Research (INPA) in Manaus, Amazonas, Brazil (catalog number: INPA 34). Etymology: The species name, bragantinus, refers to the collection site, at Bragança, in the Brazilian state of Pará, where the microparasite was discovered and described for the first time. The histological analysis shows cysts lodged internally in the gill filament, located in the intermediate portion, causing hypertrophy, a moderate increase in the size of the filaments, associated with the presence of the parasitic cysts of Myxobolus bragantinus n. sp., and a thickening of the cyst wall (Fig. 2 a–b). Under light microscopy, the ZN-stained slides revealed spores characteristic of Myxobolus (Lom & Dykova 2006), featuring polar capsules (see Fig. 2b). The location of the parasite in the gills, is known as the “intrafilamental-epithelial” (FE) type.

The third part of the catalogue of type material in the Helminthological Collection of the Oswaldo Cruz Institute/FIOCRUZ (CHIOC), comprising types deposited between 1979 and 2016, is presented to complement the first list of all types that was published in 1979. This part encompasses Acanthocephala, Nematoda and the other non-helminth phyla Cnidaria, Annelida, and Arthropoda. Platyhelminthes was covered in the first (Monogenoidea) and second (Rhabditophora Trematoda and Cestoda) parts of the catalogue published in September 2016 and March 2017, respectively. The present catalogue comprises type material for 116 species distributed across five phyla, nine classes, 50 families, and 80 genera. Specific names are listed systematically, followed by type host, infection site, type locality, and specimens with their collection numbers and references. Species classification and nomenclature are updated.

Henneguya sp. Type host: Sarpa salpa Linnaeus, 1758 goldline sea bream (Perciformes: Sparidae) Type localities: Mediterranean off Tunisia: Location 1: Gulf of Tunis (36 ° 45 ’N, 10 ° 15 ’E); Location 2: Bay of Bizerte (37 ° 20 ’ N, 9 ° 53 ’ E). Site of infection: Mesenteric vessels Prevalence: The overall prevalence is 1.2 % (4 / 330) (Fig. 9). At location 1, the prevalence of infection is 1.4 % (3 / 210) distributed as following, 03/ 2012: 0% (0/ 30); 04/ 2012: 10 % (3 / 30); 05/ 2012: 0% (0/ 30); 06/ 2012: 0% (0/ 30); 07/ 2012: 0% (0/ 30); 08/ 2012: 0% (0/ 30); 05/ 2013: 0% (0/ 20); 06/ 2013: 0% (0/ 10). At location 2, the prevalence of infection is 0.8 % (1 / 120) distributed as following, 03/ 2013: 0% (0/ 30); 04/ 2013: 0% (0/ 30); 05/ 2013: 3.3 % (1 / 30); 06/ 2013: 0% (0/ 30) (see Table 4). Mean intensity: 5 ± 2.5 cysts/infected fish (Fig. 10) (see Table 4). Type-material: Digitized photos of syntype spores were deposited in the parasitological collection of the Museum National d’Histoire Naturelle (MNHN), Paris, Coll. No. ZS 130. Description Vegetative stages. The parasite was found within mesenteric vessel of the host. Cysts were oval to round in shape and unequal in size with development asynchronous. They are measuring from 1.5 to 4 mm in diameter, filled with fluid containing a suspension of mature and immature spores which appear milky white by naked eye (Fig. 7 A). Spores (n = 30 fresh spores). Spores typical of the genus Henneguya. Mature spores were ovoid in side view with slightly attenuated posterior end (Fig. 7 C–G,J, 8 G) and ellipsoidal in sutural view (Fig. 7 I). Two shell valves smooth and equal in size with wide sutural ridge. Spore body measuring 15.26 ± 1.34 (12.6–16.2) µm in length, 9.94 ± 0.71 (9–10.8) µm in width and 8.6 ± 0.37 (8.2–9) µm in thickness. Two polar capsules were pyriform and equal in size, 5.55 ± 0.20 (5.4–5.78) Μm in length and 2.55 ± 0.1 (2.48–2.65) µm in width (n = 30). The polar filament coiled with five to six turns. Two capsulogenic nuclei located between the polar capsules (Fig. 7 F). A binucleate sporoplasm situated directly behind two polar capsules, filling almost the entire spore cavity. Ten sutural markings were distinct and usually arranged all around the circumference of the spore and being more spaced in the posterior part than at the anterior one (Figs. 7 D–E, G). Two caudal projections were filiform and long extending from posterior of spore with 49.32 ± 4.43 (42.3–55.8) µm in total length (Figs. 7 J–K, 8 G). Often, the fine distal portion of the appendage wrapped around the thicker part (Figs. 7 F, J, 8 G). Total length of spores was 64.58 ± 4.62 (58.5–70.2) Μm. Taxonomic affinities According to the scientific papers, only one histozoic species belong to genus of Hennguya Thélohan, 1892, H. neapolitana Parisi, 1912 has been described infecting the connective tissue of the renal tubule of kidney of S. salpa. The comparison between both species provides that current species differs from H. neapolitana not only by the organ host but also by the shape and the morphometric measurements (see Table 6). According to the study of Parisi (1912), H. neapolitana found in the connective tissue of the renal tubule of kidney of S. salpa with small cyst (40 to 50 µm Ø) while present species is found in the mesenteric vessels and forming a big white cysts (1-4 mm Ø) (Fig. 7 A). Concerning the shape of the spores, H. neapolitana is more ovoid than the recent form and it polar capsules often cross each other while those of present finding are almost parallel to each other and to the sutural rim. Moreover, the spores of the current species are more longer than those of H. neapolitana in length and in width of the spore body and especially in the length of caudal projections (see Table 6). Among all the species of Henneguya found in the Mediterranean Sea, no species found in the mesentery vessels of its host. All the Henneguya spp. described from the sparids are found especially infecting the heart tissue (arterial bulb), the kidney or the gills. From these known species, the recent isolate shows some similar morphological appearances to H. mbourensis and H. yoffensis Kpatcha, Faye, Diebakate, Fall & Toguebaye, 1997 infecting the kidney of Dentex canariensis (Steindachner, 1881) and the gills and heart of P. caeruleostictus (Valenciennes, 1830) respectively from Senegal, H. pagri Yokoyama, Itoh & Tanaka, 2005 infecting the bulbus arteriosus of Pagrus major (Temminck & Schlegel, 1843) from Japan, H. mauritaniensis Khlifa, Miller, Adlard, Faye & Sasal, 2012 from the arterial bulb of Pagrus caeruleostictus off Mauritania and Henneguya sp. Bahri, Benhassine & Marques, 1996 found in the gills of Sparus aurata (Linnaeus, 1758) from Tunisia that was being the same species observed by Caffara, Marcer, Florio, Quaglio & Fioravanti, 2003 in the bulbus arteriosus and the gills of the same host from Italian fish farm (see Table 6). Although all these species are very similar in shape to current species, they each exhibit one or more distinguishing characteristics. Generally all these species are shorter in total length and have spores smaller in length and width compared to those of present form (see Table 6). According to the synopsis of Henneguya (Eiras 2002, Eiras & Adriano 2012), only 2 species of Henneguya found infecting the mesentery of their hosts H. visceralis Jakovska & Nigrelli, 1953 found in Electrophorus electricus (Linnaeus, 1766) from Brazil and H. schakletoni Brickle, Kalavati & MacKenzie, 2006 in Eleginops maclovinus (Valenciennes, 1830) from Off the Falkland Islands (see Table 6). The recent species differs from H. visceralis by having a larger spores with shorter polar capsules and a very long caudal appendages. Although, most of the morphometric measurements between present finding and H. schakletoni overlap, the spores of this latter are morphologically quite different. Furthermore, the current species has larger polar capsules and longer caudal projections than those of H. schakletoni. According to the paper of Brickle et al. (2006) the spores of H. schakletoni contained one large vacuole (2.0– 3.0 Ø) placed between the posterior ends of the polar capsules and the sporoplasm which is lacking at the present species. Moreover, the cysts of our species are much larger than those of H. schekletoni (1-4 mm vs 0.5-0.8 mm) and is, therefore, a distinct species. Another species H. lateolabracis Yokoyama, Kawakami, Yasuda & Tanaka, 2003 found infecting the heart (bulb arteriosus) of the sea bass fish Lateolabrax sp. This species shows great superficial similarities in shape to our species. However, no measurement range overlap between both species (see Table 6). The recent isolate seems to have spores more bigger with longer caudal appendages compared to the spores of H. lateolabracis. Furthermore, the difference in site of infection confirms that both species are dissimilar. In light of these differences with closely related species, host organ and locality records, the myxosporean under study is considered as a different species and is reported by the first time infecting the sparid S. salpa in the Mediterranean Sea. Ecological notes In this study, Henneguya sp. found with very weak overall prevalence 1.2 %. This myxosporean has a parasitic status as scarce species. In Gulf of Tunis, the infection was only observed in April with prevalence 10 % and mean intensity 6 cysts per infected individual host whereas in Bay of Bizerte infection by this parasite was noted only in May with prevalence 3.3 % and mean intensity 4 cysts per infected individual host (see Table 4). Species Host(s) Locality Spore Polarcapsule PA(°) SL ST PCL PCW . arcuata Kalavati & Mackenzie Pagellus bogaraveo France (Monaco), Italy 6.8 ± 0.9 36.2 ± 2.7 3.7 ± 0.7 3.0 ± 0.2 ND 1999) (6.0–9.0) (32.5 –40.0) (2.5 –5.0) (2.5 –4.0) Present study) Sarpa salpa Tunisia (Gulf of Tunis) 7.5 ± 0.4 35.6 ± 3.3 3.3 ± 0.4 3 ± 0.4 150.6 ± 4.2 (7–9) (30–40) (3–4) (2.5–3.5) (142–156) . pallida Thélohan (1895) Boops boops France (Monaco) 5 25–30 ND ND ND Sarpa salpa Present study) Sarpa salpa Tunisia (Gulf of Tunis and 7.32 ± 0.61 (6.5– 28 ± 1.5 (26–30) 2.95 ± 0.47 2.92 ± 0.39 160.9 ± 4.6 Bay of Bizerte) 8) (2.5–3.6) (2.5–3.6) (154–170) . herouardi Georgévitch (1916) Sarpa salpa France (Monaco) ND ND ND ND ND Present study) Sarpa salpa Tunisia (Gulf of Tunis and 10.5 ± 1.2 21.6 ± 1.6 3.91 ± 0.25 3.89 ± 0.27 172.5 ± 6.8 Bay of Bizerte) (8–12) (20–24) (3.5–4.5) (3.5–4.5) (165–180) . sp. 1 (Present study) Sarpa salpa Tunisia (Gulf of Tunis) 7.32 ± 0.52 67.98 ± 2.44 3.26 ± 0.23 3.26 ± 0.23 α = 35.6 ± 12.3 (6.52–7.92) (64.9–70.1) (3–3.5) (3–3.5) (28–57) . sp. 2 (Present study) Sarpa salpa Tunisia (Gulf of Tunis) 9.73 ± 0.63 40.32 ± 3.83 4.2 ± 0.2 3.51 ± 0.39 150.2 ± 2.9 (9–10.5) (35–45) (4–4.5) (3–4) (146–155) . sp. 3 (Present study) Sarpa salpa Tunisia (Bay of Bizerte) 7.4 ± 0.8 30 ± 1.8 (28–33) 3 ± 0.41 3 ± 0.41 168.5 ± 4.2 (6.5–8.5) (2.5–3.5) (2.5–3.5) (162–172) . diplodae Lubat et al. (1989) Diplodus annularis Montenegro 6 (5–7) 20 (18–22) 2.25 2 ND Katharios et al. (2007) Diplodus puntazzo Greece 6.6 ± 0.5 24.0 ± 0.8 2.7 ± 0.2 2.7 ± 0.2 ND . sparusaurati Sitja-Bobadilla et Sparus aurata Spain 5.65 ± 0.74 15.76 ± 1.01 2.79 ± 0.27 2.79 ± 0.27 ND . (1995) (4.5–7.5) (14–17.5) (2.2–3.4) (2.2–3.4) . puntazzi Alama-Bermejo et al. Diplodus puntazzo Spain 9.2 ± 0.7 29 ± 2.9 4.1 ± 0.4 4 ± 0.4 166.2 ± 7.4 2011) (8.03–10.72) (23.83–34.5) (2.95–4.77) (2.9–4.6) (146.4–179.2) . sp. Alama-Bermejo et al. Diplodus annularis Spain 9.8 ± 0.8 28.8 ± 3.7 4.1 ± 0.6 4.1 ± 1.1 164.8 ± 2 2011) (7.1–13) (21.5–32.7) (3.2–5.2) (3.1–5.1) (147– 176.2) . sp. 1 Alama-Bermejo et al. Sparus aurata Spain 5 ± 0.5 17.2 ± 3.4 2.2 ± 0.4 2.1 ± 0.3 175.2 ± 4.1 2011) (3.9–5.6) (13.1–22.5) (1.6–2.7) (1.5–2.5) (166.9–179.9) . sp. 2 Alama-Bermejo et al. Sparus aurata Spain 9.9 ± 0.6 20 ± 2.1 3.8 ± 0.3 3.8 ± 0.4 169.2 ± 6.5 2011) (8.7–11.4) (16.7–24.7) (3.2–4.5) (3.2–4.5) (155.4–178.8)

Family Myxobolidae Th��lohan, 1892 Genus Henneguya Th��lohan, 1892, Published as part of Laamiri, Sayef, 2014, New observations on Myxozoa of the goldline sea bream Sarpa salpa L. 1758 (Teleostei: Sparidae) from the Mediterranean coast of Tunisia, pp. 157-190 in Zootaxa 3887 (2) on page 173, DOI: 10.11646/zootaxa.3887.2.3, http://zenodo.org/record/229268, {"references":["Thelohan, P. (1892) Myxosporidies de la vesicule biliaire des poissons. Compte Rendu Hebdomadaire des Seances de l'Academie des Sciences, 115, 1091 - 1094."]}

Henneguya sp. Type host: Sarpa salpa Linnaeus, 1758 goldline sea bream (Perciformes: Sparidae) Type localities: Mediterranean off Tunisia: Location 1: Gulf of Tunis (36 �� 45 ���N, 10 �� 15 ���E); Location 2: Bay of Bizerte (37 �� 20 ��� N, 9 �� 53 ��� E). Site of infection: Mesenteric vessels Prevalence: The overall prevalence is 1.2 % (4 / 330) (Fig. 9). At location 1, the prevalence of infection is 1.4 % (3 / 210) distributed as following, 03/ 2012: 0% (0/ 30); 04/ 2012: 10 % (3 / 30); 05/ 2012: 0% (0/ 30); 06/ 2012: 0% (0/ 30); 07/ 2012: 0% (0/ 30); 08/ 2012: 0% (0/ 30); 05/ 2013: 0% (0/ 20); 06/ 2013: 0% (0/ 10). At location 2, the prevalence of infection is 0.8 % (1 / 120) distributed as following, 03/ 2013: 0% (0/ 30); 04/ 2013: 0% (0/ 30); 05/ 2013: 3.3 % (1 / 30); 06/ 2013: 0% (0/ 30) (see Table 4). Mean intensity: 5 �� 2.5 cysts/infected fish (Fig. 10) (see Table 4). Type-material: Digitized photos of syntype spores were deposited in the parasitological collection of the Museum National d���Histoire Naturelle (MNHN), Paris, Coll. No. ZS 130. Description Vegetative stages. The parasite was found within mesenteric vessel of the host. Cysts were oval to round in shape and unequal in size with development asynchronous. They are measuring from 1.5 to 4 mm in diameter, filled with fluid containing a suspension of mature and immature spores which appear milky white by naked eye (Fig. 7 A). Spores (n = 30 fresh spores). Spores typical of the genus Henneguya. Mature spores were ovoid in side view with slightly attenuated posterior end (Fig. 7 C���G,J, 8 G) and ellipsoidal in sutural view (Fig. 7 I). Two shell valves smooth and equal in size with wide sutural ridge. Spore body measuring 15.26 �� 1.34 (12.6���16.2) ��m in length, 9.94 �� 0.71 (9���10.8) ��m in width and 8.6 �� 0.37 (8.2���9) ��m in thickness. Two polar capsules were pyriform and equal in size, 5.55 �� 0.20 (5.4���5.78) ��m in length and 2.55 �� 0.1 (2.48���2.65) ��m in width (n = 30). The polar filament coiled with five to six turns. Two capsulogenic nuclei located between the polar capsules (Fig. 7 F). A binucleate sporoplasm situated directly behind two polar capsules, filling almost the entire spore cavity. Ten sutural markings were distinct and usually arranged all around the circumference of the spore and being more spaced in the posterior part than at the anterior one (Figs. 7 D���E, G). Two caudal projections were filiform and long extending from posterior of spore with 49.32 �� 4.43 (42.3���55.8) ��m in total length (Figs. 7 J���K, 8 G). Often, the fine distal portion of the appendage wrapped around the thicker part (Figs. 7 F, J, 8 G). Total length of spores was 64.58 �� 4.62 (58.5���70.2) ��m. Taxonomic affinities According to the scientific papers, only one histozoic species belong to genus of Hennguya Th��lohan, 1892, H. neapolitana Parisi, 1912 has been described infecting the connective tissue of the renal tubule of kidney of S. salpa. The comparison between both species provides that current species differs from H. neapolitana not only by the organ host but also by the shape and the morphometric measurements (see Table 6). According to the study of Parisi (1912), H. neapolitana found in the connective tissue of the renal tubule of kidney of S. salpa with small cyst (40 to 50 ��m ��) while present species is found in the mesenteric vessels and forming a big white cysts (1-4 mm ��) (Fig. 7 A). Concerning the shape of the spores, H. neapolitana is more ovoid than the recent form and it polar capsules often cross each other while those of present finding are almost parallel to each other and to the sutural rim. Moreover, the spores of the current species are more longer than those of H. neapolitana in length and in width of the spore body and especially in the length of caudal projections (see Table 6). Among all the species of Henneguya found in the Mediterranean Sea, no species found in the mesentery vessels of its host. All the Henneguya spp. described from the sparids are found especially infecting the heart tissue (arterial bulb), the kidney or the gills. From these known species, the recent isolate shows some similar morphological appearances to H. mbourensis and H. yoffensis Kpatcha, Faye, Diebakate, Fall & Toguebaye, 1997 infecting the kidney of Dentex canariensis (Steindachner, 1881) and the gills and heart of P. caeruleostictus (Valenciennes, 1830) respectively from Senegal, H. pagri Yokoyama, Itoh & Tanaka, 2005 infecting the bulbus arteriosus of Pagrus major (Temminck & Schlegel, 1843) from Japan, H. mauritaniensis Khlifa, Miller, Adlard, Faye & Sasal, 2012 from the arterial bulb of Pagrus caeruleostictus off Mauritania and Henneguya sp. Bahri, Benhassine & Marques, 1996 found in the gills of Sparus aurata (Linnaeus, 1758) from Tunisia that was being the same species observed by Caffara, Marcer, Florio, Quaglio & Fioravanti, 2003 in the bulbus arteriosus and the gills of the same host from Italian fish farm (see Table 6). Although all these species are very similar in shape to current species, they each exhibit one or more distinguishing characteristics. Generally all these species are shorter in total length and have spores smaller in length and width compared to those of present form (see Table 6). According to the synopsis of Henneguya (Eiras 2002, Eiras & Adriano 2012), only 2 species of Henneguya found infecting the mesentery of their hosts H. visceralis Jakovska & Nigrelli, 1953 found in Electrophorus electricus (Linnaeus, 1766) from Brazil and H. schakletoni Brickle, Kalavati & MacKenzie, 2006 in Eleginops maclovinus (Valenciennes, 1830) from Off the Falkland Islands (see Table 6). The recent species differs from H. visceralis by having a larger spores with shorter polar capsules and a very long caudal appendages. Although, most of the morphometric measurements between present finding and H. schakletoni overlap, the spores of this latter are morphologically quite different. Furthermore, the current species has larger polar capsules and longer caudal projections than those of H. schakletoni. According to the paper of Brickle et al. (2006) the spores of H. schakletoni contained one large vacuole (2.0��� 3.0 ��) placed between the posterior ends of the polar capsules and the sporoplasm which is lacking at the present species. Moreover, the cysts of our species are much larger than those of H. schekletoni (1-4 mm vs 0.5-0.8 mm) and is, therefore, a distinct species. Another species H. lateolabracis Yokoyama, Kawakami, Yasuda & Tanaka, 2003 found infecting the heart (bulb arteriosus) of the sea bass fish Lateolabrax sp. This species shows great superficial similarities in shape to our species. However, no measurement range overlap between both species (see Table 6). The recent isolate seems to have spores more bigger with longer caudal appendages compared to the spores of H. lateolabracis. Furthermore, the difference in site of infection confirms that both species are dissimilar. In light of these differences with closely related species, host organ and locality records, the myxosporean under study is considered as a different species and is reported by the first time infecting the sparid S. salpa in the Mediterranean Sea. Ecological notes In this study, Henneguya sp. found with very weak overall prevalence 1.2 %. This myxosporean has a parasitic status as scarce species. In Gulf of Tunis, the infection was only observed in April with prevalence 10 % and mean intensity 6 cysts per infected individual host whereas in Bay of Bizerte infection by this parasite was noted only in May with prevalence 3.3 % and mean intensity 4 cysts per infected individual host (see Table 4). Species Host(s) Locality Spore Polarcapsule PA(��) SL ST PCL PCW . arcuata Kalavati & Mackenzie Pagellus bogaraveo France (Monaco), Italy 6.8 �� 0.9 36.2 �� 2.7 3.7 �� 0.7 3.0 �� 0.2 ND 1999) (6.0���9.0) (32.5 ���40.0) (2.5 ���5.0) (2.5 ���4.0) Present study) Sarpa salpa Tunisia (Gulf of Tunis) 7.5 �� 0.4 35.6 �� 3.3 3.3 �� 0.4 3 �� 0.4 150.6 �� 4.2 (7���9) (30���40) (3���4) (2.5���3.5) (142���156) . pallida Th��lohan (1895) Boops boops France (Monaco) 5 25���30 ND ND ND Sarpa salpa Present study) Sarpa salpa Tunisia (Gulf of Tunis and 7.32 �� 0.61 (6.5��� 28 �� 1.5 (26���30) 2.95 �� 0.47 2.92 �� 0.39 160.9 �� 4.6 Bay of Bizerte) 8) (2.5���3.6) (2.5���3.6) (154���170) . herouardi Georg��vitch (1916) Sarpa salpa France (Monaco) ND ND ND ND ND Present study) Sarpa salpa Tunisia (Gulf of Tunis and 10.5 �� 1.2 21.6 �� 1.6 3.91 �� 0.25 3.89 �� 0.27 172.5 �� 6.8 Bay of Bizerte) (8���12) (20���24) (3.5���4.5) (3.5���4.5) (165���180) . sp. 1 (Present study) Sarpa salpa Tunisia (Gulf of Tunis) 7.32 �� 0.52 67.98 �� 2.44 3.26 �� 0.23 3.26 �� 0.23 �� = 35.6 �� 12.3 (6.52���7.92) (64.9���70.1) (3���3.5) (3���3.5) (28���57) . sp. 2 (Present study) Sarpa salpa Tunisia (Gulf of Tunis) 9.73 �� 0.63 40.32 �� 3.83 4.2 �� 0.2 3.51 �� 0.39 150.2 �� 2.9 (9���10.5) (35���45) (4���4.5) (3���4) (146���155) . sp. 3 (Present study) Sarpa salpa Tunisia (Bay of Bizerte) 7.4 �� 0.8 30 �� 1.8 (28���33) 3 �� 0.41 3 �� 0.41 168.5 �� 4.2 (6.5���8.5) (2.5���3.5) (2.5���3.5) (162���172) . diplodae Lubat et al. (1989) Diplodus annularis Montenegro 6 (5���7) 20 (18���22) 2.25 2 ND Katharios et al. (2007) Diplodus puntazzo Greece 6.6 �� 0.5 24.0 �� 0.8 2.7 �� 0.2 2.7 �� 0.2 ND . sparusaurati Sitja-Bobadilla et Sparus aurata Spain 5.65 �� 0.74 15.76 �� 1.01 2.79 �� 0.27 2.79 �� 0.27 ND . (1995) (4.5���7.5) (14���17.5) (2.2���3.4) (2.2���3.4) . puntazzi Alama-Bermejo et al. Diplodus puntazzo Spain 9.2 �� 0.7 29 �� 2.9 4.1 �� 0.4 4 �� 0.4 166.2 �� 7.4 2011) (8.03���10.72) (23.83���34.5) (2.95���4.77) (2.9���4.6) (146.4���179.2) . sp. Alama-Bermejo et al. Diplodus annularis Spain 9.8 �� 0.8 28.8 �� 3.7 4.1 �� 0.6 4.1 �� 1.1 164.8 �� 2 2011) (7.1���13) (21.5���32.7) (3.2���5.2) (3.1���5.1) (147��� 176.2) . sp. 1 Alama-Bermejo et al. Sparus aurata Spain 5 �� 0.5 17.2 �� 3.4 2.2 �� 0.4 2.1 �� 0.3 175.2 �� 4.1 2011) (3.9���5.6) (13.1���22.5) (1.6���2.7) (1.5���2.5) (166.9���179.9) . sp. 2 Alama-Bermejo et al. Sparus aurata Spain 9.9 �� 0.6 20 �� 2.1 3.8 �� 0.3 3.8 �� 0.4 169.2 �� 6.5 2011) (8.7���11.4) (16.7���24.7) (3.2���4.5) (3.2���4.5) (155.4���178.8), Published as part of Laamiri, Sayef, 2014, New observations on Myxozoa of the goldline sea bream Sarpa salpa L. 1758 (Teleostei: Sparidae) from the Mediterranean coast of Tunisia, pp. 157-190 in Zootaxa 3887 (2) on pages 173-179, DOI: 10.11646/zootaxa.3887.2.3, http://zenodo.org/record/229268, {"references":["Thelohan, P. (1892) Myxosporidies de la vesicule biliaire des poissons. Compte Rendu Hebdomadaire des Seances de l'Academie des Sciences, 115, 1091 - 1094.","Parisi, B. (1912) Primo contributo alla distribusione geografica dei missosporidi in Italia. Atti della Societa Italiana de Scienza Naturalle, 50, 283 - 290.","Kpatcha, T. K., Faye, N., Diebakate, C., Fall, M. & Toguebaye, B. S. (1997) Nouvelles especes d' Henneguya Thelohan, 1895 (Myxozoa, Myxosporea) parasites des poissons marins du Senegal: Etude en microscopie photonique et electronique. Annales des Sciences Naturelles, Zoologie, 18, 81 - 91.","Yokoyama, H., Itoh, N. & Tanaka, S. (2005) Henneguya pagri n. sp. (Myxozoa: Myxosporea) causing cardiac henneguyosis in red sea bream, Pagrus major (Temminck & Schegel). Journal of Fish Diseases, 8, 479 - 487. http: // dx. doi. org / 10.1111 / j. 1365 - 2761.2005.00655. x","Khlifa, S., Miller, T. L., Adlard, R. D., Faye, N. & Sasal, P. (2012) Henneguya mauritaniensis n. sp. (Myxozoa) from the arterial bulb of Pagrus caeruleostictus (Valenciennes, 1830) off Mauritania. Parasitology Research, 111, 1287 - 1294. http: // dx. doi. org / 10.1007 / s 00436 - 012 - 2963 - 1","Bahri, S., Hassine, O. K. B. & Marques, A. (1996) Henneguya sp. (Myxosporea, Bivalvulida) infecting the gills of wild gilthead sea bream Sparus aurata L., from the coast of Tunisia. 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Laamiri, Sayef (2014): New observations on Myxozoa of the goldline sea bream Sarpa salpa L. 1758 (Teleostei: Sparidae) from the Mediterranean coast of Tunisia. Zootaxa 3887 (2): 157-190, DOI: http://dx.doi.org/10.11646/zootaxa.3887.2.3