Enrico Schwabe

The Continuing Debate on Deep Molluscan Phylogeny: Evidence for Serialia (Mollusca, Monoplacophora + Polyplacophora)

Molluscs are a diverse animal phylum with a formidable fossil record. Although there is little doubt about the monophyly of the eight extant classes, relationships between these groups are controversial. We analysed a comprehensive multilocus molecular data set for molluscs, the first to include multiple species from all classes, including five monoplacophorans in both extant families. Our analyses of five markers resolve two major clades: the first includes gastropods and bivalves sister to Serialia (monoplacophorans and chitons), and the second comprises scaphopods sister to aplacophorans and cephalopods. Traditional groupings such as Testaria, Aculifera, and Conchifera are rejected by our data with significant Approximately Unbiased (AU) test values. A new molecular clock indicates that molluscs had a terminal Precambrian origin with rapid divergence of all eight extant classes in the Cambrian. The recovery of Serialia as a derived, Late Cambrian clade is potentially in line with the stratigraphic chronology of morphologically heterogeneous early mollusc fossils. Serialia is in conflict with traditional molluscan classifications and recent phylogenomic data. Yet our hypothesis, as others from molecular data, implies frequent molluscan shell and body transformations by heterochronic shifts in development and multiple convergent adaptations, leading to the variable shells and body plans in extant lineages.

A scanning-electron microscopic study of Dickdellia labioflecta (Dell, 1990) (Gastropoda, Littorinoidea) on Colossendeis megalonyx megalonyx Fry and Hedgpeth, 1969 (Pycnogonida, Colossendeidae): a test for ectoparasitism

A scanning-electron microscopic (SEM) study of a clutch of eggs and juveniles of Dickdellia labioflecta on Colossendeis megalonyx megalonyx revealed that the young snails remove the upper layers of the pycnogonid cuticle, most probably by rasping. This way holes are created in the cuticle that could serve as a potential source of food for the snail. Measurements and statistics show that the holes have the same density as the cuticular glands. These glands are spread all over the pycnogonid cuticle. Additionally, they are filled with cytoplasmic material having a fine structure and cuticular surrounding that is typical for those glands. Hence, it is suggested that the snails get access to the interior of the pycnogonid through the holes and glands. Holes in the cuticle are absent under smaller Dickdellia specimens, but they are formed consecutively as they become older and grow larger. It is suggested that Dickdellia snails are ectoparasites because this would allow them to remain on pycnogonids until they exceed the initial egg volume, which would be difficult to explain without additional food uptake from the pycnogonid host.

Abyssal Solenogastres (Mollusca, Aplacophora) from the Northwest Pacific: Scratching the Surface of Deep-Sea Diversity Using Integrative Taxonomy

Solenogastres (Aplacophora) is a small clade of marine, shell-less worm-molluscs with close to 300 valid species. Their distribution ranges across all oceans, and whereas the vast majority of species has been collected and described from the continental shelf and slope, only few species are known from depths below 4000 m. Following traditional taxonomy, identification of specimens to species level is complex and time-consuming and requires detailed investigations of morphology and anatomy – often resulting in the exclusion of the clade in biodiversity or biogeographic studies. During the KuramBio expedition (Kuril-Kamchatka Biodiversity Studies) to the abyssal plain of the Northwest Pacific and the Kuril-Kamchatka trench, 33 solenogaster specimens were sampled from 4830 m to 5397m. Within this study we present an efficient workflow to address solenogaster diversity, even when confronted with a high degree of singletons and minute body sizes, hampering the use of single individuals for multiple morphological and molecular approaches. We combine analyses of external characters and scleritome with molecular barcoding based on a self-designed solenogaster specific set of mitochondrial primers. Overall we were able to delineate at least 19 solenogaster lineages and identify 15 species to family level and beyond. Based on our approach we identified three key lineages from the two regionally most species-rich families (Acanthomeniidae and Pruvotinidae) for deeper taxonomic investigations and describe the novel abyssal species Amboherpia abyssokurilensis sp. nov. (Cavibelonia, Acanthomeniidae) using microanatomical 3D-reconstructions. Our study more than doubles the previous records of solenogaster species from the Northwest Pacific and its marginal seas. Almost all lineages are reported for the first time from the region of the (Northwest) Pacific, vastly expanding distribution ranges of the respective clades. Moreover it doubles the number of Solenogastres collected from abyssal depths on a global scale and underlines the lack of exploratory α-diversity work in the abyssal zone for reliable species estimates in marine biodiversity.