Phylogeny and morphological evolution of the so-called bougainvilliids (Hydrozoa, Hydroidolina)

We present phylogenetic analyses (parsimony, maximum likelihood and Bayesian inference) for 69 lineages of anthoathecate hydroids based on 18 morphological characters (12 proposed for the first time) plus mitochondrial (16S and COI) and nuclear (18S and 28S) molecular markers. This study aims to test the monophyly of the present concept of the family Bougainvilliidae, assessing its phylogenetic position within Hydroidolina. Our working hypothesis is used as a context for inferring the evolution of certain morphological characters, focusing on the exoskeleton. Our results shed light on some phylogenetic uncertainties within Hydroidolina, delimiting eight well-supported linages, viz. Hydroidolina, Siphonophorae, Leptothecata, Aplanulata, Filifera II, Filifera III, Capitata and Pseudothecata taxon novum, the latter supported by four morphological synapomorphies. The monophyly of several families was not supported, viz. Bougainvilliidae, Cordylophoridae, Oceaniidae, Rathkeidae and Pandeidae. Some of the genera typically considered in Bougainvilliidae, including Bougainvillia, fell into the clade Pseudothecata, which is consistently reconstructed as the sister group of Leptothecata. We formally suggest that Dicoryne be removed from Bougainvilliidae and placed in the resurrected family Dicorynidae. The exoskeleton was a key feature in the diversification of Hydroidolina, especially with the transition from the bare hydranth to one completely enveloped within the exoskeleton. In this context, bougainvilliids exhibit several intermediate states in the development of the exosarc. Although the concatenated analysis unravels some interesting hypotheses, taxon sampling is still deficient and therefore more data are necessary for achieving a more complete understanding of the evolution and ecology of bougainvilliids and their allies.