Phylogenetic analysis of newly obtained data from the complete small subunit rDNA (18S) nuclear gene of a wide range of copepods placed the enigmatic Pectenophilus ornatus firmly in the Cyclopoida. Both maximum parsimony tree reconstruction, and Bayesian analysis operating under the GTR + I + G model of nucleotide substitution, gave identical solutions and placed P. ornatus at the base of the poecilostome families, in apposition to the mytilicolid taxa. The recently suggested assignment to the Siphonostomatoida on the basis of a tubular mouth cone in the pygmy male was rejected not only by the molecular data but also by new morphological observations. Scanning electron microscopy revealed that the appendage previously interpreted as the mandible was in reality the maxilla, the presumptive ‘labium’ only an intermaxillary outgrowth of the ventral cephalic sclerite bearing the widely separated paragnaths, and that there was no basal fusion between the labrum and the ‘posterior lip’ as in genuine siphonostomatoids. Absence of mandibles and their functional replacement by the anteriorly displaced maxillae is a unique and robust apomorphy for the Mytilicolidae and placed unequivocally P. ornatus in that family. The morphology of male Pectenophilus probably evolved as a result of global progenesis, involving early sexual maturation at the metanauplius stage and the complete cessation of somite and limb development. The molecular data were also employed to examine the relationships of two other highly modified parasitic families, the Xarifiidae (inhabiting hard corals) and the Chondracanthidae (parasitic on marine demersal fishes). Our analyses rejected the previously proposed relationship between Xarifiidae and Vahiniidae and strongly supported an Anchimolgidae + (Rhynchomolgidae + Xarifiidae) clade as sister group to the Sabelliphilidae within a monophyletic Lichomolgoidea. The obtained topology suggests that the common ancestor of this clade had already established a symbiotic relationship with scleractinian corals and that host switching occurred only secondarily in the Rhynchomolgidae, involving predominantly other cnidarian and occasionally noncnidarian hosts. Reassessment of the morphology of Parangium provided new evidence for a relationship with the xarifiids, rendering its current position in the Serpulidicolidae extremely unlikely. Both parsimony and Bayesian analyses revealed an unexpected but strongly supported relationship between the Chondracanthidae and Pseudanthessiidae. This result contrasts with earlier views advocating affinity to the Synapticolidae or Lichomolgidae, but was congruent with the previously unnoticed morphological similarity in antennary armature patterns in the first copepodid stage. The morphological grounds used to establish the Lernaeosoleidae were shown to be secondarily derived characters shared with one or several chondracanthid genera. Particularly the similarity between the Lernaeosoleidae and Markevitchielinus demonstrated that the former evolved from a mesoparasitic ancestor within the Chondracanthidae and consequently should sink as a synonym of the latter. |
