The best way to study digenean diversity combines molecular genetic methods, life-cycle studies and elaborate morphological descriptions. This approach has been barely used for one of the most widespread digenean taxa parasitizing fish – the superfamily Hemiuroidea. Here, we applied the integrative approach to the hemiuroideans from the family Derogenidae parasitizing fish at the White and Barents Seas. Analysis of 28S, 18S, 5.8S rDNA, ITS2 and cox1 gene sequences from sexually adult worms (maritae) showed genetic heterogeneity for 2 derogenid species known from this area: Derogenes varicus and Progonus muelleri. Thus, 2 pairs of genetic lineages were found: DV1 and DV2, PM1 and PM2, respectively. Data from other regions indicate that 2 more lineages of D. varicus probably exist. Based on previous records from the White and Barents Seas, we hypothesized that the cercariae found in the moonsnails (family Naticidae) belong to the Derogenidae and may help to differentiate these lineages as species. According to our results, Cercaria appendiculata from Cryptonatica affinis matched DV1, similar nameless cercariae from Euspira pallida and Amauropsis islandica matched DV2, and Cercaria octocauda from C. affinis matched PM1. We provide new data on the structure of these cercariae and discuss the life-cycle pattern of the studied digeneans.
The phylogenetic position of most xiphidiocercariae from subgroups Cercariae virgulae and Cercariae microcotylae remains unknown or unclear, even at the family level. In this paper, we studied the morphology and molecular phylogeny of 15 microcotylous and virgulate cercariae (11 new and four previously described ones). Based on morphological and molecular data, we suggested five distinct morphological types of xiphidiocercariae, which are a practical alternative to Cercariae virgulae and Cercariae microcotylae subgroups. Four of these types correspond to actual digenean taxa (Microphallidae, Lecithodendriidae, Pleurogenidae and Prosthogonimidae), while the fifth is represented by Cercaria nigrospora Wergun, 1957, which we classified on the basis of molecular data for the first time. We reassessed the relative importance of morphological characters used for the classification of virgulate and microcotylous cercariae, and discussed the main evolutionary trends within xiphidiocercariae. Now stylet cercariae can be reliably placed into several sub-taxa of Microphalloidea on the basis of their morphological features.
The Zoogonidae is the only digenean family where known cercariae lack the tail but actively search for the second intermediate host. However, the data on the zoogonid life cycles are scarce. In the present study, we elucidated and verified life cycles of the Zoogonidae from the White Sea. Using rDNA data, we showed that Pseudozoogonoides subaequiporus utilizes gastropods from the family Buccinidae as the first intermediate host and protobranch bivalves as the second one. This life cycle can be facultatively truncated: some cercariae of P. subaequiporus encyst within the daughter sporocysts. Molecular data also confirmed previous hypotheses on Zoogonoides viviapus life cycle with buccinid gastropods acting as the first intermediate hosts, and annelids and bivalves as the second intermediate hosts. We demonstrated the presence of short tail primordium in the developing cercariae of both species. Based on the reviewed and our own data, we hypothesize that the emergence of tailless cercariae in the evolution of the Zoogonidae is linked to the switch to non-arthropod second intermediate hosts, and that it possibly happened only in the subfamily Zoogoninae. Basally branching zoogonids have retained the ancestral second intermediate host and might have also retained the tail.
Few digeneans of the family Fellodistomidae are known from the Russian Arctic seas. The taxonomic status of these species, their life cycles and host range raised recurrent questions, some of which remain unanswered. To revise the species composition and life cycles of fellodistomids in the White Sea, we searched for them in several known and suspected hosts: wolffish, flatfishes (definitive), gastropods of the family Buccinidae (second intermediate) and protobranch bivalves (first intermediate). Species identification was based both on morphology and 28S ribosomal RNA gene sequences. We found Fellodistomum agnotum in the White Sea for the first time. Buccinum undatum was proved to be intermediate host of both F. agnotum and Fellodistomum fellis, and metacercariae of F. fellis were registered from two more buccinid species: Buccinum scalariforme and Neptunea despecta. We also found metacercariae of F. agnotum and F. fellis producing eggs in the second intermediate host. Two fellodistomids were found in protobranch bivalves: sporocysts and cercariae of Steringophorus furciger in Nuculana pernula, and sporocysts with large furcocercous cercariae in Ennucula tenuis. The latter were identified as F. agnotum by molecular analysis; thus, the entire life cycle of this species was reconstructed.
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