How and why diverse sexual systems evolve are fascinating evolutionary questions, but few empirical studies have dealt with these questions in animals. Pedunculate (gooseneck) barnacles show such diversity, including simultaneous hermaphroditism, coexistence of dwarf males and hermaphrodites (androdioecy), and coexistence of dwarf males and females (dioecy). Here, we report the first phylogenetically controlled test of the hypothesis that the ultimate cause of the diverse sexual systems and presence of dwarf males in this group is limited mating opportunities for non-dwarf individuals, owing to mating in small groups. Within the pedunculate barnacle phylogeny, dwarf males and females have evolved repeatedly. Females are more likely to evolve in androdioecious than hermaphroditic populations, suggesting that evolution of dwarf males has preceded that of females in pedunculates. Both dwarf males and females are associated with a higher proportion of solitary individuals in the population, corroborating the hypothesis that limited mating opportunities have favoured evolution of these diverse sexual systems, which have puzzled biologists since Darwin.

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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.

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Crabs of the genus Macrophthalmus are known to exhibit highly developed and diverse social behaviour, such as allocleaning, fighting and waving display behaviour, the first being observed widely throughout the genus. Fighting behaviour between males has been classified previously into grasping fighting and claw-extending fighting, and male waving display into four patterns, the vertical non-forward-pointing type, vertical forward-pointing type, lateral nonforward-pointing type and lateral forward-pointing type, on the basis of interspecific behaviour comparisons. To understand the evolutionary pathways of these social behavioural activities , 978-bp nucleotide sequences from mitochondrial 16S rRNA genes of 21 species, including two outgroup taxa, were analysed and a molecular phylogeny was reconstructed. The resultant tree demonstrated striking inconsistencies with the relationships inferred from morphological features. Species with similar habitat conditions showed similar morphological features, although they were not phylogenetically close relatives. Phylogenetic analysis of allocleaning behaviour suggested that it evolved once in the early history of the lineage. The analysis of fighting behaviour demonstrated that species with clawextending fighting, being a more complex behaviour than grasping fighting, are found in the most ancestral part of the phylogeny. The analysis also revealed that claw-extending fighting has evolved secondarily on two occasions, suggesting that fighting behaviour is not characterized by sufficient phylogenetic components. The superimposition of a waving pattern on to the phylogeny indicated that the lateral non-forward-pointing type has evolved from the vertical non-forward-pointing type, the lateral forward-pointing type having evolved from the vertical forward-pointing type. This scenario also appeared reasonable with respect to the behavioural trends of cheliped movements in waving.