Hundreds of genetic barcodes of the species-rich hydroid superfamily Plumularioidea (Cnidaria, Medusozoa) provide a guide toward more reliable taxonomy

Marine hydroids are important benthic components of shallow and deep waters worldwide, but their taxonomy is controversial because diagnostic morphological characters to categorize taxa are limited. Their genetic relationships are also little investigated. We tested taxonomic hypotheses within the highly speciose superfamily Plumularioidea by integrating a classical morphological approach with DNA barcoding of the 16S and COI mitochondrial markers for 659 and 196 specimens of Plumularioidea, respectively. Adding Genbank sequences, we inferred systematic relationships among 1,114 plumularioids, corresponding to 123 nominal species and 17 novel morphospecies in five families of Plumularioidea. We found considerable inconsistencies in the systematics of nominal families, genera and species. The families Kirchenpaueriidae and Plumulariidae were polyphyletic and the Halopterididae paraphyletic. Most genera of Plumularioidea are not monophyletic. Species diversity is considerably underestimated. Within our study, at least 10% of the morphologically-distinctive morphospecies are undescribed, and about 40% of the overall species richness is represented by cryptic species. Convergent evolution and morphological plasticity therefore blur systematic relationships. Additionally, cryptic taxa occur frequently in sympatry or parapatry, complicating correspondence with type material of described species. Sometimes conspecificity of different morphotypes was found. The taxonomy of hydroids requires continued comprehensive revision.