Heterochrony, generic distinction and phylogeny in the family Hydractiniidae (Hydrozoa: Cnidaria)

The taxonomy of Hydractinia, Stylactaria and Podocoryna is discussed and the three genera are merged into Hydractinia since their diagnostic characters are liable to lead to polyphyly and paraphyly, due to repeated episodes of medusa reduction via heterochrony (paedomorphosis). The phylogeny of the Hydractiniidae is reconstructed by using two outgroups, Clava and Cytaeis, both having some characters in common with the Hydractiniidae. The resulting phylogenetic trees agree in identifying affinities among Hydractinia, Kinetocodium and Hydrocorella, all with polymorphic colonies with gastrozooids having oral tentacles. The position of Clavactinia (characterized by gastrozooids with widely scattered tentacles) is at the root of the tree if Clava is the outgroup, whereas it becomes apical when the outgroup is Cytaeis. The pattern of medusa suppression is different in the two cladograms, since the presence of a medusa is a plesiomorphic feature when Cytaeis is the outgroup, whereas it becomes apomorphic when the outgroup is Clava. These inconveniences are difficult to accommodate, since medusa suppression has occurred many times in the evolution of the hydroidomedusae, and Recent species do not witness past paedomorphic events of medusa reduction properly, so that many intermediate states are probably missing.