The origin of king crabs: hermit crab ancestry under the magnifying glass

The origin of king crabs from a hermit crab ancestor has caused controversy for more than a century. While the phylogenetic position of Lithodidae within the hermit crab family Paguridae has been strengthened in recent years, several key questions regarding the evolution of lithodid crabs have remained unanswered. In particular, it has been debated which hermit crabs constitute the closest extant relatives to lithodid crabs within Paguridae. Also, the relationships of the two lithodid subfamilies, Lithodinae and Hapalogastrinae, are unresolved. Answers to these questions are crucial to the understanding of the origin of king crabs, in particular which factors were the driving forces behind leaving a protective housing, transforming to a crab-like morphology and finally developing a large body size. To address these questions, we constructed the most comprehensive molecular phylogeny of Paguridae and Lithodidae to date. Our analyses revealed a species-rich clade of hermit crabs as closest relatives to lithodid crabs within Paguridae. Hermit crabs included in this clade have a predominantly shallow-water distribution in the North Pacific, agreeing with a proposed origin of lithodid crabs in this region. We suggest that the advances resulting from abandoning a shell-inhabiting lifestyle, rather than constraints of such shelters, played a central role in carcinization in this taxon. Phylogenetic relationships within Lithodidae revealed its two subfamilies to be non-monophyletic. Small-sized, shallow-water taxa are basal in the phylogenetic tree, while an increase in size and subsequent deep-sea distribution occurred later in the evolution of the group.