Molecular phylogeny and diversity of sea pens (Cnidaria: Octocorallia: Pennatulacea) with a focus on shallow water species of the northwestern Pacific Ocean

Abstract

The order Pennatulacea, commonly known as sea pens, are colony-forming benthos belonging within subclass Octocorallia (Anthozoa, Cnidaria). Sea pens are found worldwide from shallow to deep waters, and they are important components in sandy and muddy environments. Thus far, there has been only one molecular study focusing on the phylogenetic relationships within the order Pennatulacea, which mainly treated deep-sea species, and thus information on shallow water species is still lacking. On a regional scale, the diversity of sea pens in the northwestern Pacific, including Japan and Palau, has not been well investigated. In this research, we aimed to: (1) more accurately resolve the phylogenetic relationships of sea pens with the inclusion of shallow water species, and (2) obtain a better understanding of the diversity of sea pens in Japan and Palau. Specimens were collected by SCUBA and dredging from the Ryukyu Islands in southern Japan, and from mainland Japan and Palau, and identified to at least the genus level by their morphological traits. Construction of phylogenetic trees with concatenated sequences including the mitochondrial mutS-like protein DNA mismatch repair gene mtMutS and the NADH dehydrogenase subunit 2 ND2 region were performed. The p-distances of mtMutS were calculated for estimation of species number following McFadden et al. (2011). Molecular data for 12 families and 20 genera of sea pens were used in this study. This most comprehensive study including shallow water taxa provided us with more knowledge of phylogenetic relationships. The resulting phylogenetic trees showed a topology distinguished by four large clades (clades 1–4). Families Veretillidae and Echinoptilidae are shown as not the earliest-diverging taxa. Virgulariidae and Scleroptilidae are shown as polyphyletic groups, and our results reconfirm that families Pennatulidae, Kophobelemnidae and Umbellulidae are not monophyletic groups. Overall, we collected and examined an estimated 18 species from the Ryukyu Islands, 16 species from mainland Japan, and five species from Palau. Some of these specimens represented new records from Ryukyu Islands and Palau. Previous records of these sea pens did not exist likely due to a lack of diversity research in sandy and muddy areas. These results demonstrate that many sea pens discoveries likely remain in shallow waters of the Pacific.