Relicanthus daphneae (formerly Boloceroides daphneae) was first described in 2006 as a giant sea anemone based on morphology. In 2014, its classification was challenged based on molecular data: using five genes, Relicanthus was resolved sister to zoanthideans, but with mixed support. To better understand the evolutionary relationship of Relicanthus with other early-branching metazoans, we present 15 newly-sequenced sea anemone mitochondrial genomes and a mitogenome-based phylogeny including all major cnidarian groups, sponges, and placozoans. Our phylogenetic reconstruction reveals a moderately supported sister relationship between Relicanthus and the Actiniaria. Morphologically, the cnidae of Relicanthus has apical flaps, the only existing synapomorphy for sea anemones. Based on both molecular and morphological results, we propose a third suborder (Helenmonae) within the Actiniaria to accommodate Relicanthus. Although Relicanthus shares the same gene order and content with other available actiniarian mitogenomes, it is clearly distinct at the nucleotide level from anemones within the existing suborders. The phylogenetic position of Relicanthus could reflect its association with the periphery of isolated hydrothermal vents, which, although patchy and ephemeral, harbor unique chemosynthetic communities that provide a relatively stable food source to higher trophic levels over long evolutionary timescales. The ability to colonize the deep sea and the periphery of new vent systems may be facilitated by Relicanthus’ large and extremely yolky eggs.

Hexacorals are important components of macrobenthic communities in the Southern Ocean, dominating Antarctic continental shelves. Most of the 119 sea anemones recorded for the Southern Ocean are endemic (81% and 25% endemic species and genera, respectively, one endemic family) with only two species extending beyond the limits of the Southern Ocean. Over 70% of the 83 genera in the Southern Ocean are monotypic, including half of the generic diversity in superfamily Actinostoloidea, which suggests that Antarctica has been isolated long enough for the evolution of new genera but not for many families to evolve. Here, we describe Chitinactis marmara gen. & sp. nov., a new monotypic actinostoloidean genus from Antarctica diagnosed by its unique bi-layered cuticle on column, hexamerous symmetry, unequal development of younger mesenteries and mesogleal tentacle musculature. We also re-describe and extend the geographic distribution of Scytophorus striatus, another endemic Antarctic species. Based on morphological and molecular data, we establish the phylogenetic position of C. marmara and discuss the implications of the phylogenetic position of S. striatus for the resurrection and circumscription of the family Halcampoididae and the evolution of burrowing sea anemones. Finally, we discuss evidence for an actinostoloidean deep-sea, polar lineage of burrowing sea anemones.