New Family of Allomorphic Jellyfishes, Drymonematidae (Scyphozoa, Discomedusae), Emphasizes Evolution in the Functional Morphology and Trophic Ecology of Gelatinous Zooplankton

Molecular analyses have revealed many cryptic species in the oceans, often permitting small morphological differences to be recognized as diagnosing species, but less commonly leading to consideration of cryptic ecology. Here, based on analyses of three nuclear DNA sequence markers (ribosomal 18S, 28S, and internal transcribed spacer 1 [ITS1]), two mitochondrial DNA markers (cytochrome c oxidase subunit I [COI] and ribosomal 16S), and 55 morphological features, we revise the classification of the enigmatic jellyfish genus Drymonema. We describe a new scyphozoan family, Drymonematidae, elevating the previous subfamily Drymonemidae to accommodate three species: the type species D. dalmatinum from the Mediterranean region, for which we identify a neotype; the western South Atlantic species D. gorgo; and a new species, D. larsoni from the western Atlantic and Caribbean, which also is described here. This revision emphasizes the remarkable morphological disparity of Drymonematidae from all other scyphomedusae, including allometric growth of the bell margin distal of the rhopalia, an annular zone of tentacles on the subumbrella, and ontogenetic loss of gastric filaments. Anatomical innovations are likely functionally related to predatory specialization on large gelatinous zooplankton, most notably the phylogenetically younger moon jellyfish Aurelia, indicating evolution of the feeding niche in Drymonematidae. This family-level revision contributes to the growing body of evidence that scyphomedusae are far more taxonomically rich, their biogeography is a more detailed mosaic, and their phenotypes are more nuanced than traditionally thought. Ecological and evolutionary responses to environmental change, past or future, are likely to be commensurately diverse.