Structure, Function and Evolution of Somatic Musculature in Dasydytidae (paucitubulatina, Gastrotricha)

The musculature of two species of the gastrotrich taxon Dasydytidae, Dasydytes (Dasydytes) goniathrix and Haltidytes crassus, was investigated and described using phalloidin staining, confocal microscopy and computer-aided three-dimensional data analysis. Dasydytidae is a peculiar taxon of freshwater Gastrotricha, containing species that are characterized by different adaptations to a semiplanktonic lifestyle, a rather uncommon feature among primarily benthic Gastrotricha. Like other dasydytid species studied so far, D. goniathrix and H. crassus possess a system of movable cuticular spines with an associated system of somatic oblique and segmented lateral muscles. The presence of other somatic (dorsodermal muscles R1 and R2) and visceral muscles (musculi ventrales, m. ventrolaterales, m. dorsales, m. helicoidales) known from a wide range of gastrotrich species was confirmed. Regarded from a functional perspective, the earlier proposed antagonistic role of oblique muscles (as spine abductors) and segmented lateral muscles (as adductors) is questioned for the species studied herein. Alternatively, our structural and behavioral observations suggest that muscular spine abduction in D. goniathrix is brought about by synergistic contraction of the musculi obliqua and m. laterales, and a passive adduction due to muscle relaxation and elastic recoil of the trunk and cuticle. For H. crassus, we hypothesize active muscular spine abduction by contraction of the musculi obliqua plus the last segment of m. laterales accompanied by severe cuticle deformations close to the spine insertions. Adduction is achieved by cuticle reformation due to elasticity and increase in tissue pressure brought about by muscle action, possibly of enforced dorsodermal muscles. The newly obtained and published muscular data of further gastrotrich species were gathered in a species-character matrix. Based on this data set, a maximum parsimony analysis of representatives of the Dasydytidae has been conducted. According to this analysis, there are three well-supported monophyletic lineages within likewise monophyletic Dasydytidae. The first lineage comprises the taxa Anacanthoderma, Stylochaeta and Chitonodytes, the second comprises Dasydytes, Setopus and Ornamentula, and the third represents the taxon Haltidytes. Relationships between these clades could be resolved but are only weakly supported. The new phylogenetic hypothesis is used to reconstruct the ancestral character pattern and to infer possible evolutionary transformations within the Dasydytidae.