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Among crabs of the family Ocypodidae, Ilyoplax has been known to exhibit unique mud-using territorial behavior against neighbors, including neighbor burrow plugging, barricade building, and fence building. To assess the evolution of current behavioral forms observed in Ilyoplax, 1,416-bp nucleotide sequences from the mitochondrial 12S rRNA to 16S rRNA genes of 20 species, representing four recognized subfamilies of Ocypodidae, were analyzed. The resultant phylogenetic tree revealed the subfamily Dotillinae, including Ilyoplax, to be monophyletic, with a sister group relationship with subfamily Camptandriinae. These two subfamilies were branched after Ocypodinae, with Macrophthalminae being most basal. Species of Ilyoplax fell into three different Dotillinae lineages, indicating the genus to be polyphyletic. Crabs in two of the three lineages showed differential geographic distribution and body size. Phylogenetic analyses of behavioral characters demonstrated that mud-using techniques had evolved multiple times and sequentially. From their behavioral similarity and evolutionary occurrence, fence building is hypothesized to have evolved from barricade building, and the latter, from burrow plugging. This scenario also appeared reasonable with respect to behavioral trends observed in the field. The evolution of such territorial behavior is considered to be associated with ecological conditions such as burrow fidelity and substrate condition.

Molecular phylogeny and evolution of unique mud-using territorial behavior in ocypodid crabs (Crustacea: Brachyura: Ocypodidae)