AIMS: As the most abundant cell population in the blood, erythrocytes represent an attractive source of nutrients and a protective niche to a number of pathogens. Previously, we observed the attachment of the myxozoan parasite Sphaerospora molnari to erythrocytes of its host, common carp (Cyprinus carpio), raising a number of questions about the nature of this interaction. METHODS AND RESULTS: We elucidated the impact of S molnari on the number of erythrocytes in healthy and immunocompromised fish, over a period of 6 weeks. While we observed only a mild decrease in RBC numbers in healthy individuals, we witnessed gradual and finally severe haemolytic anaemia in immunosuppressed fish. Accompanying this overt loss was increased erythropoiesis as represented by an increase of erythroblasts in the blood. In vitro, we demonstrated the uptake of host proteins from CFSE-labelled erythrocytes, ultimately inducing death of host RBCs, likely for nutrient gain of the parasite. Nevertheless, the results do not exclude a possible role of erythrocyte-derived proteins in immune evasion. CONCLUSION: Overall, the obtained data provide first evidence for the previously unknown appetite of myxozoan parasites for host erythrocytes and create an important framework for future investigations into the molecular mechanisms underlining this interaction.

• Klíčová slova
• Myxozoa, common carp, erythrocytes, flow cytometry, haemolytic anaemia, hematophagy,
• MeSH
• erytrocyty parazitologie   MeSH
• erytropoéza fyziologie   MeSH
• fylogeneze MeSH
• hemolytické anemie parazitologie   MeSH
• kapři parazitologie   MeSH
• Myxozoa fyziologie   MeSH
• nemoci ryb parazitologie   MeSH
• stravovací zvyklosti fyziologie   MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH

BACKGROUND: Parasites employ proteases to evade host immune systems, feed and replicate and are often the target of anti-parasite strategies to disrupt these interactions. Myxozoans are obligate cnidarian parasites, alternating between invertebrate and fish hosts. Their genes are highly divergent from other metazoans, and available genomic and transcriptomic datasets are limited. Some myxozoans are important aquaculture pathogens such as Sphaerospora molnari replicating in the blood of farmed carp before reaching the gills for sporogenesis and transmission. Proliferative stages cause a massive systemic lymphocyte response and the disruption of the gill epithelia by spore-forming stages leads to respiratory problems and mortalities. In the absence of a S. molnari genome, we utilized a de novo approach to assemble the first transcriptome of proliferative myxozoan stages to identify S. molnari proteases that are upregulated during the first stages of infection when the parasite multiplies massively, rather than in late spore-forming plasmodia. Furthermore, a subset of orthologs was used to characterize 3D structures and putative druggable targets. RESULTS: An assembled and host filtered transcriptome containing 9436 proteins, mapping to 29,560 contigs was mined for protease virulence factors and revealed that cysteine proteases were most common (38%), at a higher percentage than other myxozoans or cnidarians (25-30%). Two cathepsin Ls that were found upregulated in spore-forming stages with a presenilin like aspartic protease and a dipeptidyl peptidase. We also identified downregulated proteases in the spore-forming development when compared with proliferative stages including an astacin metallopeptidase and lipases (qPCR). In total, 235 transcripts were identified as putative proteases using a MEROPS database. In silico analysis of highly transcribed cathepsins revealed potential drug targets within this data set that should be prioritised for development. CONCLUSIONS: In silico surveys for proteins are essential in drug discovery and understanding host-parasite interactions in non-model systems. The present study of S. molnari's protease arsenal reveals previously unknown proteases potentially used for host exploitation and immune evasion. The pioneering dataset serves as a model for myxozoan virulence research, which is of particular importance as myxozoan diseases have recently been shown to emerge and expand geographically, due to climate change.

• Klíčová slova
• Aquaculture, Drug targets, In silico screening, Myxozoa, Parasite, Proteases,
• MeSH
• antiparazitární látky farmakologie   terapeutické užití   MeSH
• faktory virulence MeSH
• fylogeneze MeSH
• kapři mikrobiologie   MeSH
• Myxozoa genetika   růst a vývoj   MeSH
• nemoci ryb parazitologie   terapie   MeSH
• objevování léků MeSH
• parazitární nemoci u zvířat parazitologie   terapie   MeSH
• proteasy genetika   MeSH
• transkriptom MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• Názvy látek
• antiparazitární látky MeSH
• faktory virulence MeSH
• proteasy MeSH

It is assumed that complex life cycles in cnidarian parasites belonging to the Myxozoa result from incorporation of vertebrates into simple life cycles exploiting aquatic invertebrates. However, nothing is known about the driving forces and implementation of this event, though it fostered massive diversification. We performed a comprehensive search for myxozoans in evolutionary ancient fishes (Chondrichthyes), and more than doubled existing 18S rDNA sequence data, discovering seven independent phylogenetic lineages. We performed cophylogenetic and character mapping methods in the largest monophyletic dataset and demonstrate that host and parasite phylogenies are strongly correlated, and that tectonic changes may explain phylogeographic clustering in recent skates and softnose skates, in the Atlantic. The most basal lineages of myxozoans inhabit the bile of chondrichthyans, an immunologically privileged site and protective niche, easily accessible from the gut via the bile duct. We hypothesize that feed-integration is a likely mechanism of host acquisition, an idea supported by feeding habits of chimaeras and ancient sharks and by multiple entries of different parasite lineages from invertebrates into the new host group. We provide exciting first insights into the early evolutionary history of ancient metazoan parasites in a host group that embodies more evolutionary distinctiveness than most other vertebrates.

• Klíčová slova
• Chondrichthyes, cnidaria, co-diversification, co-phylogeny, feed-integration, migration, myxozoa, phylogeography,
• Publikační typ
• časopisecké články MeSH

BACKGROUND: Sphaerospora molnari is a myxozoan parasite causing skin and gill sphaerosporosis in common carp (Cyprinus carpio) in central Europe. For most myxozoans, little is known about the early development and the expansion of the infection in the fish host, prior to spore formation. A major reason for this lack of information is the absence of laboratory model organisms, whose life-cycle stages are available throughout the year. RESULTS: We have established a laboratory infection model for early proliferative stages of myxozoans, based on separation and intraperitoneal injection of motile and dividing S. molnari stages isolated from the blood of carp. In the present study we characterize the kinetics of the presporogonic development of S. molnari, while analyzing cellular host responses, cytokine and systemic immunoglobulin expression, over a 63-day period. Our study shows activation of innate immune responses followed by B cell-mediated immune responses. We observed rapid parasite efflux from the peritoneal cavity (< 40 hours), an initial covert infection period with a moderate proinflammatory response for about 1-2 weeks, followed by a period of parasite multiplication in the blood which peaked at 28 days post-infection (dpi) and was associated with a massive lymphocyte response. Our data further revealed a switch to a massive anti-inflammatory response (up to 1456-fold expression of il-10), a strong increase in the expression of IgM transcripts and increased number of IgM+ B lymphocytes, which produce specific antibodies for the elimination of most of the parasites from the fish at 35 dpi. However, despite the presence of these antibodies, S. molnari invades the liver 42 dpi, where an increase in parasite cell number and indistinguishable outer cell membranes are indicative of effective exploitation and disguise mechanisms. From 49 dpi onwards, the acute infection changes to a chronic one, with low parasite numbers remaining in the fish. CONCLUSIONS: To our knowledge, this is the first time myxozoan early development and immune modulation mechanisms have been analyzed along with innate and adaptive immune responses of its fish host, in a controlled laboratory system. Our study adds important information on host-parasite interaction and co-evolutionary adaptation of early metazoans (Cnidaria) with basic vertebrate (fish) immune systems and the evolution of host adaptation and parasite immune evasion strategies.

• Klíčová slova
• B cells, Cyprinus carpio, Cytokines, Host–parasite interaction, IgM, Myxozoa, Sphaerospora molnari, Teleost,
• MeSH
• buněčná imunita MeSH
• cytokiny metabolismus   MeSH
• hlavová ledvina metabolismus   MeSH
• humorální imunita MeSH
• interakce hostitele a parazita MeSH
• kapři imunologie   parazitologie   MeSH
• modely nemocí na zvířatech MeSH
• Myxozoa růst a vývoj   imunologie   MeSH
• nemoci ryb imunologie   parazitologie   MeSH
• parazitární nemoci u zvířat imunologie   parazitologie   MeSH
• spory MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• Názvy látek
• cytokiny MeSH

During a survey on the myxosporean fauna of Rajiformes from the Atlantic coast of Argentina, in waters off Buenos Aires Province (34°-42°S; 53°-62°W), the gall bladders of 217 specimens belonging to seven species of skates, representatives of two families, were examined. As a result, three species of Chloromyxum Mingazzini, 1890, namely C. atlantoraji n. sp., C. zearaji n. sp. and C. riorajum Azevedo, Casal, Garcia, Matos, Teles-Grilo and Matos, 2009 were found infecting three endemic host species, the spotback skate Atlantoraja castelnaui (Arhynchobatidae), the yellownose skate Zearaja chilensis (Rajidae) and the Rio skate Rioraja agassizii (Arhynchobatidae), respectively. These species were described based on myxospore morphology and morphometry characterization, as well as by providing their small subunit ribosomal DNA (SSU rDNA) sequences. The SSU rDNA-based phylogenetic analyses showed that these three species constituted a well-established monophyletic subclade within the marine Chloromyxum clade, while branches subtending the other Chloromyxum species were poorly resolved or unresolved, independently of the host taxonomic identities (Carchariniformes, Myliobatiformes, Orectolobiformes, Pristiophoriformes, Rajiformes, Squaliformes and Torpediniformes) and/or host geographic distribution (Atlantic coast of Portugal, Atlantic coast of the USA, Australian waters or Mediterranean Sea). The possible causes of these discrepancies are discussed, providing new insights into the phylogeny of the marine Chloromyxum clade.

Lors d’une étude de la faune des Myxozoaires des Rajiformes de la côte atlantique argentine, dans les eaux situées au large de la province de Buenos Aires (34°–42°S; 53°–62°O), les vésicules biliaires de 217 spécimens appartenant à sept espèces, représentants deux familles, ont été examinés. En conséquence, trois espèces de Chloromyxum Mingazzini, 1890, à savoir C. atlantoraji n. sp., C. zearaji n. sp. et C. riorajum Azevedo, Casal, Garcia, Matos, Teles-Grilo et Matos, 2009 ont été trouvées, infectant trois espèces hôtes endémiques, Atlantoraja castelnaui (Arhynchobatidae), Zearaja chilensis (Rajidae) et Rioraja agassizii (Arhynchobatidae), respectivement. Ces espèces sont décrites sur la base de la morphologie et de la morphométrie des myxospores, ainsi qu’en fournissant leurs petites séquences d’ADN ribosomal (SSU ADNr). Les analyses phylogénétiques basées sur l’ADNr SSU ont montré que ces trois espèces constituaient un sous-clade monophylétique bien établi dans le clade des Chloromyxum marins, tandis que les branches sous-jacentes aux autres espèces de Chloromyxum étaient mal ou non résolues, indépendamment des identités taxonomiques hôtes (Carchariniformes, Myliobatiformes, Orectolobiformes, Pristiophoriformes, Rajiformes, Squaliformes et Torpediniformes) et/ou de la répartition géographique de l’hôte (côte atlantique du Portugal, côte atlantique des États-Unis, eaux australiennes ou mer Méditerranée). Les causes possibles de ces divergences sont discutées, fournissant de nouvelles informations sur la phylogénie du clade des Chloromyxum marins.

• MeSH
• druhová specificita MeSH
• fylogeneze * MeSH
• mikroskopie elektronová rastrovací MeSH
• Myxozoa klasifikace   genetika   izolace a purifikace   MeSH
• nemoci ryb epidemiologie   parazitologie   MeSH
• oceány a moře MeSH
• parazitární nemoci u zvířat epidemiologie   parazitologie   MeSH
• rejnokovití parazitologie   MeSH
• ribozomální DNA genetika   MeSH
• sekvence nukleotidů MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• Geografické názvy
• Argentina epidemiologie   MeSH
• oceány a moře MeSH
• Názvy látek
• ribozomální DNA MeSH

BACKGROUND: Myxozoa are extremely diverse microscopic parasites belonging to the Cnidaria. Their life-cycles alternate between vertebrate and invertebrate hosts, predominantly in aquatic habitats. Members of the phylogenetically well-defined Sphaerospora (sensu stricto) clade predominantly infect the urinary system of marine and freshwater fishes and amphibians. Sphaerosporids are extraordinary due to their extremely long and unique insertions in the variable regions of their 18S and 28S rDNA genes and due to the formation of motile proliferative stages in the hosts' blood. To date, DNA sequences of only 19 species have been obtained and information on the patterns responsible for their phylogenetic clustering is limited. METHODS: We screened 549 fish kidney samples from fish of various geographical locations, mainly in central Europe, to investigate sphaerosporid biodiversity microscopically and by 18S rDNA sequences. We performed multiple phylogenetic analyses to explore phylogenetic relationships and evolutionary trends within the Sphaerospora (s.s.) clade, by matching host and habitat features to the resultant 18S rDNA trees. The apparent co-clustering of species from related fish hosts inspired us to further investigate host-parasite co-diversification, using tree-based (CoRE-PA) and distance-based (ParaFit) methods. RESULTS: Our study considerably increased the number of 18S rDNA sequence data for Sphaerospora (s.s.) by sequencing 17 new taxa. Eight new species are described and one species (Sphaerospora diminuta Li & Desser, 1985) is redescribed, accompanied by sufficient morphological data. Phylogenetic analyses showed that sphaerosporids cluster according to their vertebrate host order and habitat, but not according to geography. Cophylogenetic analyses revealed a significant congruence between the phylogenetic trees of sphaerosporids and of their vertebrate hosts and identified Cypriniformes as a host group of multiple parasite lineages and with high parasite diversity. CONCLUSIONS: This study significantly contributed to our knowledge of the biodiversity and evolutionary history of the members of the Sphaerospora (s.s.) clade. The presence of two separate phylogenetic lineages likely indicates independent historical host entries, and the remarkable overlap of the larger clade with vertebrate phylogeny suggests important coevolutionary adaptations. Hyperdiversification of sphaerosporids in cypriniform hosts, which have undergone considerable radiations themselves, points to host-driven diversification.

• Klíčová slova
• Host-parasite codiversification, Myxozoa, Phylogeny, Sphaerospora sensu stricto, Taxonomy, Teleost,
• MeSH
• biodiverzita * MeSH
• biologická evoluce MeSH
• Cnidaria MeSH
• fylogeneze * MeSH
• Myxozoa klasifikace   genetika   izolace a purifikace   fyziologie   MeSH
• nemoci ryb parazitologie   MeSH
• parazitární nemoci u zvířat genetika   parazitologie   MeSH
• ribozomální DNA genetika   MeSH
• ryby klasifikace   genetika   parazitologie   MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• Názvy látek
• ribozomální DNA MeSH

INTRODUCTION: Intertidal rock pools where fish and invertebrates are in constant close contact due to limited space and water level fluctuations represent ideal conditions to promote life cycles in parasites using these two alternate hosts and to study speciation processes that could contribute to understanding the roles of parasitic species in such ecosystems. MATERIAL AND METHODS: Gall bladder and liver samples from five clinid fish species (Blenniiformes: Clinidae) were morphologically and molecularly examined to determine the diversity, prevalence, distribution and host specificity of Ceratomyxa parasites (Cnidaria: Myxozoa) in intertidal habitats along the coast of South Africa. Phylogenetic relationships of clinid ceratomyxids based on the SSU rDNA, LSU rDNA and ITS regions were assessed additionally to the investigation of population genetic structure of Ceratomyxa cottoidii and subsequent comparison with the data known from type fish host Clinus cottoides. RESULTS AND DISCUSSION: Seven Ceratomyxa species including previously described Ceratomyxa dehoopi and C. cottoidii were recognized in clinids. They represent a diverse group of rapidly evolving, closely related species with a remarkably high prevalence in their hosts, little host specificity and frequent concurrent infections, most probably as a result of parasite radiation after multiple speciation events triggered by limited host dispersal within restricted spaces. C. cottoidii represents the most common clinid parasite with a population structure characterized by young expanding populations in the south west and south east coast and by older populations in equilibrium on the west coast of its distribution. Parasite and fish host population structures show overlapping patterns and are very likely affected by similar oceanographic barriers possibly due to reduced host dispersal enhancing parasite community differentiation. While fish host specificity had little impact on parasite population structure, the habitat preference of the alternate invertebrate host as well as tidal water exchange may be additional crucial variables affecting the dispersal and associated population structure of C. cottoidii.

• MeSH
• fylogeneze MeSH
• interakce hostitele a parazita genetika   fyziologie   MeSH
• Myxozoa klasifikace   genetika   MeSH
• nemoci ryb parazitologie   MeSH
• parazitární nemoci u zvířat parazitologie   MeSH
• paraziti klasifikace   genetika   MeSH
• populační genetika metody   MeSH
• ribozomální DNA genetika   MeSH
• ryby parazitologie   MeSH
• stadia vývoje genetika   MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• práce podpořená grantem MeSH
• Geografické názvy
• Jihoafrická republika MeSH
• Názvy látek
• ribozomální DNA MeSH

BACKGROUND: Wildlife repatriation represents an opportunity for parasites. Reintroduced hosts are expected to accumulate generalist parasites via spillover from reservoir hosts, whereas colonization with specialist parasites is unlikely. We address the question of how myxozoan parasites, which are characterized by a complex life-cycle alternating between annelids and fish, can invade a reintroduced fish species and determine the impact of a de novo invasion on parasite diversity. We investigated the case of the anadromous allis shad, Alosa alosa (L.), which was reintroduced into the Rhine approximately 70 years after its extinction in this river system. METHODS: We studied parasites belonging to the Myxozoa (Cnidaria) in 196 allis shad from (i) established populations in the French rivers Garonne and Dordogne and (ii) repatriated populations in the Rhine, by screening the first adults returning to spawn in 2014. Following microscopical detection of myxozoan infections general myxozoan primers were used for SSU rDNA amplification and sequencing. Phylogenetic analyses were performed and cloned sequences were analyzed from individuals of different water sources to better understand the diversity and population structure of myxozoan isolates in long-term coexisting vs recently established host-parasite systems. RESULTS: We describe Hoferellus alosae n. sp. from the renal tubules of allis shad by use of morphological and molecular methods. A species-specific PCR assay determined that the prevalence of H. alosae n. sp. is 100 % in sexually mature fish in the Garonne/Dordogne river systems and 22 % in the first mature shad returning to spawn in the Rhine. The diversity of SSU rDNA clones of the parasite was up to four times higher in the Rhine and lacked a site-specific signature of SNPs such as in the French rivers. A second myxozoan, Ortholinea sp., was detected exclusively in allis shad from the Rhine. CONCLUSIONS: Our data demonstrate that the de novo establishment of myxozoan infections in rivers is slow but of great genetic diversity, which can only be explained by the introduction of spores from genetically diverse sources, predominantly via straying fish or by migratory piscivorous birds. Long-term studies will show if and how the high diversity of a de novo introduction of host-specific myxozoans succeeds into the establishment of a local successful strain in vertebrate and invertebrate hosts.

• Klíčová slova
• Alosa alosa, Diversity, Hoferellus alosae n. sp., Host reintroduction, Myxozoa, Parasite population structure, SNPs,
• MeSH
• fylogeneze MeSH
• genetická variace * MeSH
• jednonukleotidový polymorfismus MeSH
• Myxozoa genetika   izolace a purifikace   MeSH
• nemoci ryb epidemiologie   parazitologie   MeSH
• parazitární nemoci u zvířat parazitologie   MeSH
• pohyb MeSH
• řeky parazitologie   MeSH
• ribozomální DNA MeSH
• ryby parazitologie   MeSH
• stadia vývoje MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• práce podpořená grantem MeSH
• Geografické názvy
• Francie epidemiologie   MeSH
• Názvy látek
• ribozomální DNA MeSH

BACKGROUND: Swim bladder inflammation (SBI) is an important disease of common carp fingerlings in Central Europe. In the 1980s, its etiology was ascribed to multicellular proliferative stages of the myxozoan parasite Sphaerospora dykovae (formerly S. renicola). S. dykovae was reported to proliferate in the blood and in the swim bladder prior to the invasion of the kidney, where sporogony takes place. Due to the presence of emerging numbers of proliferative myxozoan blood stages at different carp culture sites in recent years we analysed cases of SBI, for the first time, using molecular diagnostics, to identify the myxozoan parasites present in diseased swim bladders. METHODS: We amplified myxozoan SSU rDNA in a non-specific approach and compared the species composition in swim bladders at culture sites where carp demonstrated 1. No signs of SBI, 2. Minor pathological changes, and 3. Heavy SBI. Based on DNA sequences, we determined the localisation and distribution of the most frequent species by in situ hybridisation, thereby determining which myxozoans are involved in SBI. RESULTS: Large multicellular myxozoan swim bladder stages characterised heavy SBI cases and were identified as S. dykovae, however, blood stages were predominantly represented by Sphaerospora molnari, whose numbers were greatly increased in carp with mild and heavy SBI, compared with SBI-free fish. S. molnari was found to invade different organs and cause inflammatory changes also in the absence of S. dykovae. One site with mild SBI cases was characterised by Buddenbrockia sp. infection in different organs and a general granulomatous response. CONCLUSIONS: We provide evidence that the etiology of SBI can vary in relation to culture site and disease severity and that emerging numbers of S. molnari in the blood represent an important co-factor or precondition for SBI.

• MeSH
• časové faktory MeSH
• kapři MeSH
• klonování DNA MeSH
• Myxozoa klasifikace   genetika   MeSH
• nemoci ryb parazitologie   patologie   MeSH
• parazitární nemoci u zvířat parazitologie   patologie   MeSH
• prevalence MeSH
• roční období MeSH
• vzdušné vaky parazitologie   patologie   MeSH
• zánět parazitologie   patologie   veterinární   MeSH
• zvířata MeSH
• Check Tag
• zvířata MeSH
• Publikační typ
• časopisecké články MeSH
• práce podpořená grantem MeSH

UNLABELLED: Myxosporeans (Myxozoa), microscopic metazoan parasitic organisms, are poorly studied in the Arctic region. Our survey of benthic and pelagic fish (n = 234) collected in Isfjorden (Svalbard, Norway) together with detailed morphological and molecular examination revealed the presence of nine myxosporean species. We compared observed myxosporean diversity with diversity documented in regions close to the Arctic and revealed that water depth rather than geographic distribution is an important factor influencing myxosporean fauna. WE DESCRIBE THREE NEW MYXOSPOREAN SPECIES: Zschokkella siegfriedi n. sp. from kidney of Boreogadus saida, Parvicapsula petuniae n. sp. from the urinary bladder of Gymnocanthus tricuspis, and Sinuolinea arctica n. sp. from the urinary bladder of Myoxocephalus scorpius. We characterise Latyspora-like organism from kidney of Clupea harengus. We provide new data for Ceratomyxa porrecta, Myxidium gadi, Myxidium finnmarchicum, Schulmania aenigmatosa, and Parvicapsula irregularis comb. nov. The phylogenetic analyses including the newly obtained SSU and LSU rDNA data revealed that most of the species studied cluster in the marine urinary clade within the marine myxosporean lineage. Newly obtained sequences including the first molecular data for the member of the genus Schulmania, substantially enriched the Zschokkella subclade. C. porrecta and the two Myxidium species cluster within the Ceratomyxa and marine Myxidium clade, respectively. Newly described species, Z. siegfriedi n. sp., was revealed to be morphologically indistinguishable but genetically diverse from Zschokkella hildae known from numerous gadid fish. Therefore, we consider Z. siegfriedi to be a cryptic myxosporean species that might be misidentified with Z. hildae. A Latyspora-like organism was found to be taxonomically problematic due to its suture line and its distant phylogenetic position from the type species Latyspora scomberomori did not allow us to assign it to the genus Latyspora. Based on an increased taxon sampling and SSU + LSU rDNA-based phylogeny, evolutionary trends within the marine urinary clade are investigated.

• Klíčová slova
• Arctic, Marine urinary clade, Myxosporea, Phylogeny, Schulmania, Zschokkella,
• Publikační typ
• časopisecké články MeSH