Cryptic diversity, characterized by morphologically similar but genetically distinct species, poses significant challenges to traditional taxonomic methods. Within monogeneans parasitizing northwest African barbels, this complexity hampers species identification, limiting our understanding of diversity, distribution and evolutionary relationships. Supported by previously published genetic data, we morphologically delineate herein 9 Gyrodactylus species from Morocco. Newly described species include G. agnesei sp. nov. and G. benhoussai sp. nov. from Luciobarbus rabatensis, with the latter also found on Carasobarbus fritschii, and both G. deburonae sp. nov. and G. marruecosi sp. nov. from L. massaensis. Additionally, G. diakini sp. nov. and G. louiziae sp. nov. were identified from L. rifensis and L. yahyaouii, respectively. Pterocapoeta maroccana harboured G. pterocapoetai sp. nov., morphologically resembling G. shigoleyae sp. nov. from sympatric L. zayanensis. We also examined taxonomical discrepancies between Gyrodactylus species from L. ksibi and L. pallaryi, evaluated the status of previously described G. nyingiae and described G. qninbai sp. nov. from L. ksibi. Our findings highlight the conservative morphology in northwest African Gyrodactylus, characterized by an ancestral median ridge in the ventral bar membrane, similar to that found in species from Eurasia. Subtle phenotypic features, like bifurcations in dorsal bars and proportions of marginal hooks, serve as diagnostic traits. We further evidenced a potential host-switching event from northwest African to Iberian hosts, correlating with the region's geological history and cyprinid dispersal events during intermittent closures of the Strait of Gibraltar. These insights illuminate the complex evolutionary processes driving gyrodactylid diversification in the West Mediterranean.

• Keywords
• Morocco, Southwest Mediterranean, cyprinids, gyrodactylids, monogenea,
• MeSH
• Cyprinidae * parasitology   MeSH
• Phylogeny MeSH
• Trematode Infections * veterinary   parasitology   epidemiology   MeSH
• Fish Diseases * parasitology   epidemiology   MeSH
• Trematoda * classification   anatomy & histology   genetics   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Geographicals
• Morocco epidemiology   MeSH

Monogenea (Platyhelminthes), mainly gill and fin ectoparasites of fish, are often recognized as host specific and morphologically and ecologically diverse. These parasites exhibit high species diversity at the level of host species or individual fish specimens. Using case studies, especially those widely performed in Dactylogyrus parasitizing cyprinoid fish, this article presents current knowledge on the ecology and evolution of congeneric gill monogeneans. The important aspects of the ecology of congeneric monogeneans are highlighted, in particular: host specificity expressed at several host levels (from strict specificity to phylogenetic specificity), microhabitat specificity expressed by restricted positions on fish gills to facilitate intraspecific mating, and the link between microhabitat preference and morphological adaptation (i.e., sclerotized structures of the haptor) or reproductive isolation. From the evolutionary perspective, this study focused on the processes of the speciation and diversification of congeneric monogeneans, highlighting the role of host switch as the most prominent coevolutionary event, accompanied in some cases mostly by intrahost speciation or cospeciation, as revealed by cophylogenetic studies. Here, important knowledge on evolutionary patterns of host specificity, microhabitat specificity, and morphological adaptation is presented. Host-specific monogeneans may represent an important tool for studying the historical biogeography of their hosts. Specifically, in the case of freshwater fish hosts exhibiting disjunctive distribution, they reflect both historical and contemporary contacts. The role of host-specific congeneric monogeneans in revealing historical intercontinental and intracontinental contacts between freshwater fish is highlighted. Finally, the importance of the role of genetic coadaptation, limiting the presence of host-specific monogeneans in hybrid fish, is emphasized.

TITLE: Monogènes spécifiques à leur hôte parasitant les poissons d’eau douce : écologie et évolution des associations hôtes-parasites. ABSTRACT: Les Monogènes (Plathelminthes), principalement ectoparasites des branchies et des nageoires des poissons, sont souvent reconnus comme spécifiques à leur hôte et diversifiés morphologiquement et écologiquement. Ces parasites présentent une grande diversité d’espèces au niveau des espèces hôtes ou des spécimens individuels de poissons. À l’aide d’études de cas (en particulier celles largement réalisées chez les Dactylogyrus parasitant les poissons cyprinoïdes), les connaissances actuelles sur l’écologie et l’évolution des monogènes branchiaux congénères sont présentées. Les aspects importants de l’écologie des monogènes congénères sont mis en évidence, en particulier la spécificité à l’hôte exprimée à plusieurs niveaux de l’hôte (de la spécificité stricte à la spécificité phylogénétique), la spécificité du microhabitat exprimée par des positions restreintes sur les branchies des poissons pour faciliter l’accouplement intraspécifique, et le lien entre la préférence du microhabitat et l’adaptation morphologique (c’est-à-dire les structures sclérifiées du hapteur) ou l’isolement reproductif. Du point de vue évolutif, l’étude a été concentrée sur les processus de spéciation et de diversification des monogènes congénères, soulignant le rôle du changement d’hôte comme l’événement coévolutif le plus important, accompagné surtout dans certains cas de spéciation ou de cospéciation intra-hôte, comme le révèlent les études cophylogénétiques. Des connaissances importantes sont présentées ici sur les modèles évolutifs de spécificité d’hôte, de spécificité de microhabitat et d’adaptation morphologique. Les monogènes spécifiques à l’hôte peuvent représenter un outil important pour étudier la biogéographie historique de leurs hôtes. Plus précisément, dans le cas des poissons d’eau douce hôtes présentant une distribution disjonctive, ils reflètent à la fois des contacts historiques et contemporains. Le rôle des monogènes congénères spécifiques à leur hôte dans la révélation des contacts intercontinentaux et intracontinentaux historiques entre poissons d’eau douce est mis en évidence. Enfin, l’importance du rôle de la coadaptation génétique limitant la présence de monogènes spécifiques à l’hôte chez les poissons hybrides est soulignée.

• Keywords
• Fish parasites, Genetic coadaptation, Host specificity, Host-parasite coevolution, Monogeneans, Species coexistence,
• MeSH
• Biological Evolution * MeSH
• Species Specificity MeSH
• Ecosystem MeSH
• Phylogeny MeSH
• Host Specificity * MeSH
• Trematode Infections veterinary   parasitology   MeSH
• Host-Parasite Interactions * MeSH
• Fish Diseases * parasitology   MeSH
• Platyhelminths physiology   classification   genetics   isolation & purification   MeSH
• Fishes parasitology   MeSH
• Fresh Water * parasitology   MeSH
• Trematoda physiology   classification   genetics   MeSH
• Gills * parasitology   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Review MeSH

The freshwaters of Iraq harbour a high diversity of endemic and phylogenetically unique species. One of the most diversified fish groups in this region is cyprinoids, and although their distribution is relatively well known, their monogenean parasites have only rarely been investigated. Herein, we applied an integrative approach, combining morphology with molecular data, to assess the diversity and phylogeny of cyprinoid-associated monogenean parasites. A total of 33 monogenean species were collected and identified from 13 endemic cyprinoid species. The highest species diversity was recorded for Dactylogyrus (Dactylogyridae, 16 species) and Gyrodactylus (Gyrodactylidae, 12 species). Four species of Dactylogyrus and 12 species of Gyrodactylus were identified as new to science and described. Two other genera, Dogielius (Dactylogyridae) and Paradiplozoon (Diplozoidae), were represented only by 4 and 1 species, respectively. Phylogenetic analyses of the Dactylogyrus and Gyrodactylus species revealed that the local congeners do not form a monophyletic group and are phylogenetically closely related to species from other regions (i.e. Europe, North Africa and Eastern Asia). These findings support the assumption that the Middle East served as an important historical crossroads for the interchange of fauna between these 3 geographic regions.

• Keywords
• Cyprinoidei, Dactylogyrus, Dogielius, Gyrodactylus, Middle East, Paradiplozoon, phylogeny, species diversity,
• MeSH
• Phylogeny MeSH
• Fishes MeSH
• Trematoda * genetics   MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Geographicals
• Iraq epidemiology   MeSH
• Africa, Northern MeSH
• Middle East MeSH

Hybrid generations usually face either a heterosis advantage or a breakdown, that can be expressed by the level of parasite infection in hybrid hosts. Hybrids are less infected by parasites than parental species (especially F1 generations) or more infected than parental species (especially post-F1 generations). We performed the experiment with blood-feeding gill parasite Paradiplozoon homoion (Monogenea) infecting leuciscid species, Abramis brama and Rutilus rutilus, their F1 generation and two backcross generations. Backcross generations tended to be more parasitized than parental lines and the F1 generation. The number of differentially expressed genes (DEGs) was lower in F1 hybrids and higher in backcross hybrids when compared to each of the parental lines. The main groups of DEGs were shared among lines; however, A. brama and R. rutilus differed in some of the top gene ontology (GO) terms. DEG analyses revealed the role of heme binding and erythrocyte differentiation after infection by blood-feeding P. homoion. Two backcross generations shared some of the top GO terms, representing mostly downregulated genes associated with P. homoion infection. KEGG analysis revealed the importance of disease-associated pathways; the majority of them were shared by two backcross generations. Our study revealed the most pronounced DEGs associated with blood-feeding monogeneans in backcross hybrids, potentially (but not exclusively) explainable by hybrid breakdown. The lower DEGs reported in F1 hybrids being less parasitized than backcross hybrids is in line with the hybrid advantage.

• Keywords
• Monogenea, Paradiplozoon homoion, RNA seq, differential gene expression, freshwater fish, hybrid breakdown, hybrid heterosis, hybridization,
• Publication type
• Journal Article MeSH

BACKGROUND: Several hypotheses have been proposed to explain parasite infection in parental species and their hybrids. Hybrid heterosis is generally applied to explain the advantage for F1 generations of hybrids exhibiting a lower level of parasite infection when compared to parental species. Post-F1 generations often suffer from genetic incompatibilities potentially reflected in the higher level of parasite infection when compared to parental species. However, the presence of specific parasites in an associated host is also limited by close coevolutionary genetic host-parasite associations. This study focused on monogenean parasites closely associated with two leuciscid fish species-common bream and roach-with the aim of comparing the level of monogenean infection between parental species and hybrids representing two F1 generations with different mtDNA and two backcross generations with different cyto-nuclear compositions. RESULTS: Monogenean infection in F1 generations of hybrids was lower when compared to parental species, in line with the hybrid heterosis hypothesis. Monogenean infection in backcross generations exhibited similarities with the parental species whose genes contributed more to the backcross genotype. The distribution of monogeneans associated with one or the other parental species showed the same asymmetry with a higher proportion of roach-associated monogeneans in both F1 generations and backcross generation with roach in the paternal position. A higher proportion of common bream-associated monogeneans was found in backcross generation with common bream in the paternal position. CONCLUSIONS: Our study indicated that cyto-nuclear incompatibilities in hybrids do not induce higher monogenean infection in backcross generations when compared to parental species. However, as backcross hybrids with a higher proportion of the genes of one parental taxon also exhibited high level of this parental taxon-associated parasites, host-parasite coevolutionary interactions seem to play an obvious role in determining the level of infection of host-specific monogeneans in hybrids.

• Keywords
• Freshwater fish, Genetic incompatibilities, Host specificity, Host-parasite coevolution, Hybrid heterosis, Hybrids, Monogeneans,
• Publication type
• Journal Article MeSH

The genes of the major histocompatibility complex (MHC) are an essential component of the vertebrate immune system and MHC genotypes may determine individual susceptibility to parasite infection. In the wild, selection that favors MHC variability can create situations in which interspecies hybrids experience a survival advantage. In a wild system of two naturally hybridizing leuciscid fish, we assessed MHC IIB genetic variability and its potential relationships to hosts' ectoparasite communities. High proportions of MHC alleles and parasites were species-specific. Strong positive selection at specific MHC codons was detected in both species and hybrids. MHC allele expression in hybrids was slightly biased towards the maternal species. Controlling for a strong seasonal effect on parasite communities, we found no clear associations between host-specific parasites and MHC alleles or MHC supertypes. Hybrids shared more MHC alleles with the more MHC-diverse parental species, but expressed intermediate numbers of MHC alleles and positively selected sites. Hybrids carried significantly fewer ectoparasites than either parent species, suggesting a hybrid advantage via potential heterosis.

• MeSH
• Alleles * MeSH
• Bayes Theorem MeSH
• Biological Evolution * MeSH
• Models, Biological MeSH
• Cyprinidae genetics   MeSH
• Major Histocompatibility Complex MeSH
• Hybridization, Genetic * MeSH
• Hybrid Vigor * genetics   MeSH
• Crosses, Genetic MeSH
• Linear Models MeSH
• Selection, Genetic * MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH

BACKGROUND: Host specificity is one of the outputs of the coevolution between parasites and their associated hosts. Several scenarios have been proposed to explain the pattern of parasite distribution in parental and hybrid genotypes ranging from hybrid resistance to hybrid susceptibility. We hypothesized that host-parasite co-adaptation limits the infection of host-specific parasites in hybrid genotypes even under the condition of the high frequency of hybrids. The experimental monogenean infection in pure breeds of Blicca bjoerkna and Abramis brama and cross-breeds (the F1 generation of hybrids) under the condition of similar frequencies of pure and hybrid genotypes was investigated. We also examined the potential effect of the maternal origin of hybrids (potential co-adaptation at the level of mitochondrial genes) on monogenean abundance. METHODS: Pure breeds of two cyprinids and two cross-breeds (one with B. bjoerkna, the next with A. brama in the maternal positions) were exposed to infection by monogeneans naturally occurring in B. bjoerkna and A. brama. The experiment was run under similar frequencies of the four breed lines. RESULTS: We showed similar levels of monogenean infection in B. bjoerkna and A. brama. However, each species harboured specific monogenean fauna. Hybrids harboured all monogenean species specifically infecting one or the other species. Monogenean infection levels, especially those of Dactylogyrus specific to A. brama, were lower in hybrids. For the majority of host-specific parasites, there was no effect of the maternal origin of hybrids on monogenean abundance. Asymmetry was found in the distribution of specific parasites in favour of specialists of B. bjoerkna in the monogenean communities of hybrids. CONCLUSIONS: Our results indicate that the maternal mtDNA of hybrids is not an important predictor of host-specific monogenean infection, which may suggest that mitochondrial genes are not strongly involved in the coadaptation between monogeneans and their associated hosts. The asymmetry of species-specific parasites suggests similarity between the molecular components of the immune mechanisms in hybrids and B. bjoerkna. Our results revealed a difference between the degree of host-parasite coadaptation in specific parasites of A. brama and the degree of host-parasite coadaptation in specific parasites of B. bjoerkna and their associated hosts.

• Keywords
• Coadaptation, Cyprinid fish, Host specificity, Hybridization, Monogenean infection,
• MeSH
• Cyprinidae genetics   parasitology   MeSH
• Phylogeny MeSH
• Adaptation, Physiological MeSH
• Genotype MeSH
• Host Specificity * MeSH
• Trematode Infections * MeSH
• Host-Parasite Interactions MeSH
• Genes, Mitochondrial * MeSH
• Fish Diseases parasitology   MeSH
• Fishes MeSH
• Trematoda MeSH
• Animals MeSH
• Check Tag
• Animals MeSH
• Publication type
• Journal Article MeSH

The epicontinental fauna of the Iberian Peninsula is strongly influenced by its geographical history. As the possibilities for dispersion of organisms into and from this region were (and still are) limited, the local fauna consists almost exclusively of endemic species. Almost all Iberian freshwater fishes of the families Leuciscidae and Cyprinidae are endemic and on-going research on these taxa continually uncovers new species. Nevertheless, information on their host-specific parasites remains scarce. In this study, we investigate the diversity and phylogenetic relationships in monogeneans of the genus Dactylogyrus (gill ectoparasites specific to cyprinoid fish) in the Iberian Peninsula. Twenty-two species were collected and identified from 19 host species belonging to Cyprinidae and Leuciscidae. A high degree of endemism was observed, with 21 Dactylogyrus species reported from Iberia only and a single species, D. borealis, also reported from other European regions. Phylogenetic analysis split the endemic Iberian Dactylogyrus into two well-supported clades, the first encompassing Dactylogyrus parasitizing endemic Luciobarbus spp. only, and the second including all Dactylogyrus species of endemic leuciscids and four species of endemic cyprinids. Species delimitation analysis suggests a remarkable diversity and existence of a multitude of cryptic Dactylogyrus species parasitizing endemic leuciscids (Squalius spp. and representatives of Chondrostoma s.l.). These results suggest a rapid adaptive radiation of Dactylogyrus in this geographically isolated region, closely associated with their cyprinoid hosts. Moreover, phylogenetic analysis supports that Dactylogyrus parasites colonized the Iberian Peninsula through multiple dispersion events.

• Keywords
• Cyprinoidea, Iberian Peninsula, Monogenea, host-specificity, phylogeny, species delimitation,
• MeSH
• Biodiversity MeSH
• Adaptation, Biological MeSH
• Biological Evolution * MeSH
• Cyprinidae * MeSH
• Trematode Infections parasitology   veterinary   MeSH
• Fish Diseases parasitology   MeSH
• Trematoda physiology   MeSH
• Animals MeSH
• Check Tag
• Male MeSH
• Female MeSH
• Animals MeSH
• Publication type
• Journal Article MeSH
• Research Support, Non-U.S. Gov't MeSH
• Geographicals
• Portugal MeSH
• Spain MeSH