Phylogeny and taxonomy of Haloclavidae (Verrill, 1899) with a redescription of the parasitic, burrowing sea anemone, Peachia chilensis Carlgren, 1931

Abstract

Haloclavidae Verrill, 1899 is a family of burrowing sea anemones grouped within the superfamily Actinioidea (Rafinesque, 1815). Currently, it includes 30 species in 10 genera. Characters given for this family in descriptions of its taxa have not been consistent, with numerous exceptions to the expectations of the familial diagnosis. Previous phylogenetic analyses have shown that Haloclavidae is potentially a polyphyletic group, but resolution of relationships of the few representatives of Haloclavidae included in analyses has been problematic. Here we address questions of monophyly and affinity of Haloclavidae using three mitochondrial and two nuclear markers. We assess the monophyly of Haloclavidae in the context of all major lineages of Actiniaria Hertwig, 1882, emphasizing diversity of superfamily Actinioidea. We use parsimony-based character optimization to interpret the distribution of key traits in the superfamily. We find that Haloclavidae is not monophyletic and propose two new families, Peachiidae fam. nov. and Harenactidae fam. nov., while also retaining some species in the family Haloclavidae, so that taxonomy better reflects relationships and diversity of the group. In addition, we redescribe a species within the newly created Peachiidae, Peachia chilensis Carlgren, 1931. We use recent larval samples obtained in Antofagasta, Chile, and the histological slides from the original description to redescribe P. chilensis, to provide a complete account of cnidae, external, and internal morphology. Finally, we compare P. chilensis to other burrowing anemones found in Chile and provide an understanding of the genus Peachia that reflects recent phylogenetic perspective on diversity of anemones previously assigned to family Haloclavidae.