Dactylogyrus species (Dactylogyridae: Monogenea) are a group of monogenean gill parasites that are highly specific to freshwater fish of the family Cyprinidae. Dactylogyrus species were sampled from 19 cyprinids and one percid collected in Europe. Using partial 18S rDNA and ITS1 sequences, a phylogeny of 51 Dactylogyrus species was reconstructed to investigate the patterns of parasite speciation and diversification. Three main Dactylogyrus lineages were recognized from all phylogenetic trees, that is, analysis of 18S rDNA alone and combined 18SrDNA and ITS1. The first lineage associates the Dactylogyrus species of Cyprinus carpio and Carassius auratus of the Cyprininae; the second associates Dactylogyrus species of the Gobioninae, Pseudorasbora parva of the Rasborinae, and Ctenopharyngodon idella of the Cyprininae; and the third associates Dactylogyrus species of the Leuciscinae and Alburninae and Barbus barbus of the Cyprininae. Our results suggest that the genus Dactylogyrus is of quite recent origin and that these three lineages separated from each other in a very short period of time. Host subfamily mapping onto the parasite tree inferred from analysis of the combined dataset showed that the Cyprininae could be plesiomorphic hosts for Dactylogyrus. Dactylogyrus parasites would have secondarily colonized the Percidae and representatives of the Leuciscinae, Alburninae, Gobioninae, and Rasborinae. Comparison of host and parasite phylogenetic relationships indicated that a very high number of parasite duplications occurred within two of the three Dactylogyrus lineages. Dactylogyrus diversification can be mainly explained by sympatric intrahost speciation events that seem to be correlated to strict host specificity. Moreover, the present study shows that the congeneric parasites speciating within one host tend to occupy niches within hosts differing at least in one niche parameter.
Patterns and likely processes connected with evolution of host specificity in congeneric monogeneans parasitizing fish species of the Cyprinidae were investigated. A total of 51 Dactylogyrus species was included. We investigated (1) the link between host specificity and parasite phylogeny; (2) the morphometric correlates of host specificity, parasite body size, and variables of attachment organs important for host specificity; (3) the evolution of morphological adaptation, that is, attachment organ; (4) the determinants of host specificity following the hypothesis of specialization on more predictable resources considering maximal body size, maximal longevity, and abundance as measures of host predictability; and (5) the potential link between host specificity and parasite diversification. Host specificity, expressed as an index of host specificity including phylogenetic and taxonomic relatedness of hosts, was partially associated with parasite phylogeny, but no significant contribution of host phylogeny was found. The mapping of host specificity into the phylogenetic tree suggests that being specialist is not a derived condition for Dactylogyrus species. The different morphometric traits of the attachment apparatus seem to be selected in connection with specialization of specialist parasites and other traits favored as adaptations in generalist parasites. Parasites widespread on several host species reach higher abundance within hosts, which supports the hypothesis of ecological specialization. When separating specialists and generalists, we confirmed the hypothesis of specialization on a predictable resource; that is, specialists with larger anchors tend to live on fish species with larger body size and greater longevity, which could be also interpreted as a mechanism for optimizing morphological adaptation. We demonstrated that ecology of host species could also be recognized as an important determinant of host specificity. The mapping of morphological characters of the attachment organ onto the parasite phylogenetic tree reveals that morphological evolution of the attachment organ is connected with host specificity in the context of fish relatedness, especially at the level of host subfamilies. Finally, we did not find that host specificity leads to parasite diversification in congeneric monogeneans.
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