The description and delineation of trematode species is a major ongoing task. Across the field there has been, and currently still is, great variation in the standard of this work and in the sophistication of the proposal of taxonomic hypotheses. Although most species are relatively unambiguously distinct from their congeners, many are either morphologically very similar, including the major and rapidly growing component of cryptic species, or are highly variable morphologically despite little to no molecular variation for standard DNA markers. Here we review challenges in species delineation in the context provided to us by the historical literature, and the use of morphological, geographical, host, and molecular data. We observe that there are potential challenges associated with all these information sources. As a result, we encourage careful proposal of taxonomic hypotheses with consideration for underlying species concepts and frank acknowledgement of weaknesses or conflict in the data. It seems clear that there is no single source of data that provides a wholly reliable answer to our taxonomic challenges but that nuanced consideration of information from multiple sources (the ‘integrated approach’) provides the best possibility of developing hypotheses that will stand the test of time.
Morphological discrimination of species is problematic in many digenean taxa. Parasites of marine fish from the genus Lecithaster Lühe, 1901 are a good example of this. Our goal was to understand which species of Lecithaster infect fish in the White Sea, and reveal their life cycles. We collected specimens of maritae from nine fish species, analysed their morphology and sequenced 28S ribosomal DNA and internal transcribed spacer 2 (ITS2). Contrary to previous accounts, all of them belong to a single species, Lecithaster salmonis Yamaguti, 1934, which was previously only recorded from the Pacific. Morphologically, our maritae specimens were highly variable, sharing characters of L. salmonis, Lecithaster confusus Odhner, 1905 and Lecithaster gibbosus (Rudolphi, 1802) Lühe, 1901. This variability did not correlate with the moderate differences in ITS2 among the specimens, and neither did the fish host species. Members of the subfamily Salmoninae appear to be the best suited definitive hosts, judging from the intensity rates. The intermediate hosts were also discovered: the first is Cryptonatica affinis (Gmelin, 1791) and the second are planktonic copepods. These lifecycle data from the White Sea are consistent with L. salmonis species identification and with the distribution of this species in the North Pacific. The geographical range of L. salmonis seems to be interrupted, and we discuss possible ways of L. salmonis expansion.
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.
Studying complexes of cryptic or pseudocryptic species opens new horizons for the understanding of speciation processes, an important yet vague issue for the digeneans. We investigated a hemiuroidean trematode Lecithaster salmonis across a wide geographic range including the northern European seas (White, Barents, and Pechora), East Siberian Sea, and the Pacific Northwest (Sea of Okhotsk and Sea of Japan). The goals were to explore the genetic diversity within L. salmonis through mitochondrial (cox1 and nad5 genes) and ribosomal (ITS1, ITS2, 28S rDNA) marker sequences, to study morphometry of maritae, and to revise the life cycle data. Mitochondrial markers showed that L. salmonis is likely divided into six lineages (referred to as operational taxonomic units, OTUs), which often occur in sympatry, sometimes in a single host specimen. Variation in rDNA was not consistent with that in the mitochondrial markers. Morphometric analysis of maritae was performed for four out of six OTUs; it showed that some OTUs had significant differences from the others, but some did not. The effect of host species on the morphometric characteristics cannot be excluded. Intramolluscan stages were identified for two OTUs; they differed clearly by the structure of cercariae and also by the species of the first intermediate host. The case of L. salmonis is instructive in how different criteria for species delimitation can contradict each other. We regard this as a sign of recent or ongoing speciation and suggest using the name Lecithaster cf. salmonis. The most promising criteria to differentiate genetic lineages within L. cf. salmonis are first intermediate hosts and morphological characteristics of the cercariae: shape of the delivery tube and caudal cyst, and length of the filamentous appendage.
Mycetomorpha vancouverensis is an enigmatic parasitic barnacle from the family Mycetomorphidae, known for its unclear phylogenetic position within Rhizocephala. Specimens of M. vancouverensis were collected from infected Neocrangon communis shrimps near the South Kuril Islands. Detailed morphological studies were conducted using histological techniques and scanning electron microscopy, and 18S rDNA sequences were used to resolve the phylogenetic position of M. vancouverensis within Rhizocephala. Morphological analysis revealed a complex externa structure with spermatogenic bodies and the presence of a main trunk in the interna. Some of the side branches of the interna invade the host’s nerve tissue and are modified into specialized goblet-shaped organs. These invasive rootlets are morphologically similar to the goblet-shaped organs in the family Peltogastridae. Molecular analysis confirmed that the Mycetomorphidae is a sister group to the Peltogastridae. At the same time, peltogastrids were recovered as polyphyletic. Our findings contribute to the understanding of the evolution of parasitic barnacles and highlight the unique adaptations of Mycetomorphidae within the broader context of Rhizocephala.
20 March 2023 in Journal of Invertebrate Pathology