Structure and formation of the uncini in Pectinaria koreni, Pectinaria auricoma (Terebellida) and Spirorbis spirorbis (Sabellida): inplications for annelid phylogeny and the position of the Pogonophora

Abstract

Setation is an important taxonomic character of the Annelida. Within this taxon, Terebellida and Sabellida both have transverse rows of short, apically toothed setae which are situated inside the neuropodial rim. The apical spines are curved and their tips point anteriorly. These setae are termed uncini. In the terebellidans Pectinaria koreni, Pectinaria auricoma and in the sabellidan Spirorbis spirorbis, these setae arise from a follicle which consists of a chaetoblast and two follicle cells. The special structure of the uncini is a result of temporal modifications of the actin-filament system of the chaetoblast and changing spatial interactions between the chaetoblast and the follicle cells during the formation of these setae. Once the uncinus is formed, the microvilli are withdrawn and electron-dense material is deposited in the remaining canals. The microvilli are replaced by short processes of the chaetoblast, and the actin-filament system is replaced by a system of intermediate filaments which help to mechanically attach the uncinus to the follicle. Such uncini are also described in both pogonophoran groups, the Perviata and the Obturata (Vestimentifera). In several structural details they correspond to those of the species investigated in this paper, so that the hypothesis of a homology of the uncini seems to be justified. This hypothesis leads to the conclusion that uncini evolved in the common stem lineage of Pogonophora, Terebellida and Sabellida, implying a monophyletic origin of these three taxa. The uncini are compared to the hooked setae of the Arenicolida, Maldanida and Psammodrilida, which are also aligned in transverse rows inside the neurophodial rim. Hooked setae and uncini are hypothesized to be homologous. It, therefore, can be concluded that Arenicolida, Maldanida and Psammodrilida are closely related to the monophylum consisting of Terebellida, Sabellida and Pogonophora, and that these six taxa share a common ancestor, which evolved transverse rows of setae with apically curved spines and a formative site lateral to the edge of the neuropodial rim. According to the phylogenetic relationships proposed here, the Pogonophora are a subordinate taxon within the Annelida.