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.

The aims of the study were (1) to describe the Dactylogyrus communities living on selected South European Barbus species, (2) to analyse morphometric variability of their attachment and reproductive organs, and (3) to perform molecular phylogenetic analyses, in order to investigate the mode of speciation in Dactylogyrus species parasitizing congeneric hosts. In Bulgaria, Dactylogyrus crivellius, D. dyki and D. petenyi were found on B. balcanicus, and D. dyki on B. cyclolepis. In Spain, Dactylogyrus carpathicus and D. dyki were detected on B. meridionalis. Morphometric analyses of D. dyki revealed significant differences in the attachment and reproductive organs when individuals from different Barbus species were compared. Two monophyletic groups were recognized from the molecular phylogenetic analyses: the first included D. carpathicus and D. crivellius which have large body size and anchors, with a weakly supported basal position of D. malleus from B. barbus; the second included D. dyki and D. petenyi which have small body and anchor sizes. The comparison of host and parasite phylogenies did not indicate the intrahost speciation. Intraspecific molecular variability was found between individuals of D. dyki and D. carpathicus from different Barbus species, suggesting the need for a taxonomic revision for these species.

Phylogenetic analyses based on the partial large subunit rDNA (LSU) sequences of polyonchoinean monogeneans belonging to the Dactylogyridea and Monocotylidea were generated to investigate relationships among various subfamilies of the Dactylogyridae sensu Kritsky & Boeger, 1989. Monophyly of the Dactylogyridae was supported by all analyses performed. Status of the Ancyrocephalidae sensu Bychowsky & Nagibina, 1978 and Ancyrocephalinae sensu Kritsky & Boeger, 1989 was revised based on the present data. All phylogenetic analyses indicated polyphyletic origins of the Ancyrocephalidae and Ancyrocephalinae. Freshwater species of Ancyrocephalinae (Actinocleidus, Ancyrocephalus, Cleidodiscus and Urocleidus) and Ancylodiscoidinae (Thaparocleidus) collected from the fish in European waters were positioned at the base of the Dactylogyridae. The Dactylogyrinae formed a monophyletic group, sister to a clade including the Pseudodactylogyrinae and the tropical and subtropical Ancyrocephalinae. Analyses including only data set on Dactylogyridea were focused on relationships between representatives of the Asian and European Dactylogyrus species. Dactylogyrus species formed a monophyletic group, and the parasite colonization appeared to follow the dispersal history of the Cyprinidae from Asia to Europe. Three lineages of Dactylogyrus species were recognized: the first including species specific to hosts of Asian origin, the second by Dactylogyrus species from Chinese fish hosts, and the third included Dactylogyrus species from European cyprinids and one species from a percid host. The position of D. cryptomeres from Gobio gobio seems to be unresolved.

Aggregation is one of the distinctive features in parasite-host relationships, which has generally been studied at the level of host communities. Parasite aggregation at the infracommunity level may nevertheless be important for intraspecific interactions such as parasite mating success and opportunities for cross-fertilization. In the present paper, we studied the infracommunity aggregation of 3 highly abundant Dactylogyrus (Monogenea) species occurring on the gills of crucian carp (Carassius carassius). In line with the previous work on monogenean communities, we observed no competition between the species. At the species level, parasites were distributed unevenly on the gills showing aggregation in the majority of infracommunities. However, aggregation decreased with increasing parasite abundance, which supports the hypothesis that less aggregation may be needed to ensure successful mating when the distance to a potential mate decreases with increasing number of conspecifics. Lack of interspecific interactions, species specific site selection and the importance of study scale for aggregation patterns in dactylogyrids are discussed.