Exploring evolutionary trends within the Pennellidae (Copepoda: Siphonostomatoida) using molecular data.

The family Pennellidae comprises ecto- and mesoparasitic copepods on marine fishes. Although a preliminary scheme of phylogenetic relationships of pennellids based on morphological characters exists, it is difficult to objectively define character states because of their highly modified bodies and reduced appendages. This molecule-based study analysed phylogenetic relationships among seven genera and 12 species of pennellids, using 18S and 28S ribosomal DNA sequences in order to infer evolutionary trends within the family. Our molecular analysis recovered three clades (Clade-I, Peniculus; Clade-II, Haemobaphes-Lernaeocera-Phrixocephalus-Exopenna-Lernaeenicus radiatus; and Clade-III, Pennella-Lernaeenicus spp.). This result was congruent with some of the morphology-based phylogenetic relationships previously proposed but did not support a sister group comprising Exopenna, Phrixocephalus and Pennella. The second and third offshoots after the divergence of Clade-I species are characterized by reduced body tagmosis and changes in lifestyle from ectoparasites to mesoparasites. In some gill parasites of Clade-II, their sigmoid-shaped bodies and coiled egg strings have likely evolved in adaptation to the limited available space within the gill cavities of the hosts. Phrixocephalus is an eye parasite in Clade-II, which also has coiled egg strings, may have descended from an ancestral gill parasite. All species of Clade-III are characterized by the possession of a head region with processes deeply embedded into the host tissues and functioning as an anchor.