Anatomy of the serotonergic nervous system of an entoproct creeping-type larva and its phylogenetic implications
Although the internal phyletic relationships of Spiralia (and Lophotrochozoa) remain unresolved, recent progress has been made due to molecular phylogenetic analyses as well as developmental studies of crucial taxa such as Mollusca, Sipuncula, or Annelida. Despite this progress, the phylogenetic position of a number of phyla, such as Entoprocta, remains problematic, mainly due to their unique morphology, their aberrant mode of development, and their exclusion in most large-scale phylogenetic analyses. In order to extend the morphological dataset of this enigmatic taxon, we herein describe the anatomy of the serotonergic nervous system of the creeping-type larva of Loxosomella murmanica. The apical organ is very complex and comprises six to eight centrally positioned flask cells and eight bipolar peripheral cells. In addition, a prototroch nerve ring, an anterior nerve loop, a paired buccal nerve, and an oral nerve ring are found. Moreover, the larva of L. murmanica has one pair of pedal and one pair of lateral longitudinal nerve cords and thus expresses a tetraneurous condition. Several paired serotonergic perikarya, which form contact with the pedal nerve cords but not with the lateral ones, are found along the anterior–posterior axis. The combination of a complex larval serotonergic apical organ and (adult) tetraneury, comprising one pair of ventral and one pair of more dorsally situated lateral longitudinal nerve cords without ganglia, has so far only been reported for basal molluscs and may be diagnostic for a mollusc–entoproct clade. In addition, the larva of Loxosomella expresses a mosaic of certain neural features that are also found in other larval or adult Spiralia, e.g., a prototroch nerve ring, an anterior nerve loop, and a buccal nervous system.