Evaluation of Some Vulval Appendages in Nematode Taxonomy

A survey of the nature and phylogenetic distribution of nematode vulval appendages revealed 3 major classes based on composition, position, and orientation that included membranes, flaps, and epiptygmata. Minor classes included cuticular inflations, protruding vulvar appendages of extruded gonadal tissues, vulval ridges, and peri-vulval pits. Vulval membranes were found in Mermithida, Triplonchida, Chromadorida, Rhabditidae, Panagrolaimidae, Tylenchida, and Trichostrongylidae. Vulval flaps were found in Desmodoroidea, Mermithida, Oxyuroidea, Tylenchida, Rhabditida, and Trichostrongyloidea. Epiptygmata were present within Aphelenchida, Tylenchida, Rhabditida, including the diverged Steinernematidae, and Enoplida. Within the Rhabditida, vulval ridges occurred in Cervidellus, peri-vulval pits in Strongyloides, cuticular inflations in Trichostrongylidae, and vulval cuticular sacs in Myolaimus and Deleyia. Vulval membranes have been confused with persistent copulatory sacs deposited by males, and some putative appendages may be artifactual. Vulval appendages occurred almost exclusively in commensal or parasitic nematode taxa. Appendages were discussed based on their relative taxonomic reliability, ecological associations, and distribution in the context of recent 18S ribosomal DNA molecular phylogenetic trees for the nematodes. Characters were found to be distributed across subsets of terminal and phylogenetically distant taxa, demonstrating considerable homoplasy. Accurate definitions, terminology, and documentation of the taxonomic distribution of vulval appendages are important in evaluations of hypotheses for either parallelism and developmental constraint or convergence and adaptation.