From commensalism to parasitism within a genus-level clade of barnacles.

Understanding how animals evolve to become parasites is key to unravelling how biodiversity is generated as a whole, as parasites could account for half of all species richness. Two significant impediments to this are that parasites fossilize poorly and that they retain few clear shared morphological features with non-parasitic relatives. Barnacles include some of the most astonishingly adapted parasites with the adult body reduced to just a network of tubes plus an external reproductive body, but how they originated from the sessile, filter-feeding form is still a mystery. Here, we present compelling molecular evidence that the exceedingly rare scale-worm parasite barnacle Rhizolepas is positioned within a clade comprising species currently assigned to Octolasmis, a genus exclusively commensal with at least six different phyla of animals. Our results imply that species in this genus-level clade represent an array of species at various transitional stages from free-living to parasitic in terms of plate reduction and host-parasite intimacy. Diverging only about 19.15 million years ago, the route to parasitism in Rhizolepas was associated with rapid modifications in anatomy, a pattern that was likely true for many other parasitic lineages.

Molecular and Morphological Assessment of Juvenile and Adult Forms in the Giant Worm Eunice Cf. Aphroditois (Annelida: Eunicidae) and Its Phylogenetic Position in the Family.

Eunice aphroditois (Pallas, 1788) is a large polychaete worm (up to 3 m in length) and the type species of the genus. In Japan, a similar but potentially different species, Eunice cf. aphroditois, is distributed mainly in the rocky shores of the temperate and warm Pacific coasts. Juveniles and adults were suggested to be distinguished by their body color. The juvenile form was previously regarded as distinct species, Eunice flavopicta Izuka, 1912 and Eunice ovalifera Fauvel, 1936, although they are now considered synonymous with E. aphroditois. In this study, we revisited the validity of the present taxonomy based on morphological observations including SEM and microCT, and three molecular markers (cytochrome c oxidase subunit I [COI], 16S rRNA, and histone H3 genes) and investigated the phylogenetic position of E. cf. aphroditois in the family Eunicidae using the combined dataset of three genes (COI + 16S rRNA + 18S rRNA). The adult and juvenile forms were different in body size, color, the distribution of the branchiae and subacicular hooks, and maxillae shape, but not in other characteristics. One individual showed an intermediate body color between the two forms. The adult and juvenile forms shared major haplotypes and the maximum K2P genetic distance of COI was 1.7%, which can be considered within intraspecific variation. In the phylogenetic tree based on the combined gene dataset, E. cf. aphroditois was closely related to Eunice roussaei Quatrefages, 1866 and Eunice cf. violaceomaculata Ehlers, 1887, which are large species from the Mediterranean Sea and the Caribbean Sea, respectively.

Recent origin and diversification accompanied by repeated host shifts of thallus-mining flies (Diptera: Agromyzidae) on liverworts and hornworts.

Despite the vast diversity of phytophagous insects that feed on vascular plants (tracheophytes), insects that feed on bryophytes remain understudied. Agromyzidae, one of the most species-rich phytophagous clades in Diptera, consists mainly of leaf-mining species that feed on tracheophytes. However, a recent discovery of thallus-mining species on liverworts and hornworts within the Liriomyza group of Phytomyzinae provides an opportunity to study host shifts between tracheophytes and bryophytes. This study aimed to explore the origin and diversification of thallus-miners and estimate the pattern and timing of host shifts. Phylogenetic analysis of Phytomyzinae has revealed that the thallus-mining agromyzids formed a separate clade, which was sister to a fern pinnule-miner. The diversification of bryophyte-associated agromyzids since the Oligocene involved multiple host shifts across various bryophyte taxa. The diversification of the thallus-mining Phytoliriomyza may have occurred at the same time as the leaf-mining agromyzid flies on herbaceous plants, indicating a dynamic history of interactions between bryophytes and herbivores in angiosperms-dominated ecosystems.

Diversity underfoot of agromyzids (Agromyzidae, Diptera) mining thalli of liverworts and hornworts.

Agromyzidae is a dipteran family that has diversified as internal plant feeders. Although most agromyzid species feed on herbaceous angiosperms, only a limited number of species has been recorded as miners of bryophytes. Extensive searches and rearing of bryophytivores in the Japanese Archipelago were made, resulting in that thallus-mining agromyzids are overwhelmingly widespread and diverse on thalloid liverworts and hornworts. By examining the morphology of adult flies, it was revealed that the agromyzid fauna comprise 39 species, of which 37 species are newly described. All the species are assigned to the genus Phytoliriomyza Hendel based on some shared morphological character states as follows: costa reaching M1; orbital setulae minute and erect (rarely proclinate); male epandrium with combs of fused tubercle-like setae and/or hypertrophied arms bearing tubercle-like setae; male distiphallus comprising a pair of stout, extended tubules; female cercus with two stout, apical, trichoid sensilla. Of the 39 agromyzid species in Japan, 36 species are associated with liverworts: 5 spp. on Marchantia (Marchantiaceae), 2 spp. on Dumortiera (Dumortieraceae), 3 spp. on Plagiochasma, 1 sp. on Asterella, 6 spp. on Reboulia (Aytoniaceae), 1 sp. on Wiesnerella (Wiesnerellaceae), 15 spp. on Conocephalum (Conocephalaceae), and 3 spp. on Riccia (Ricciaceae). Three species are associated with hornworts: 1 sp. on Folioceros (Anthocerotaceae), 1 sp. on Megaceros (Dendrocerotaceae), and 1 sp. on Notothylas,Phaeoceros (Notothyladaceae), and Anthoceros (Anthocerotaceae). The results suggest that 37 of the 39 species are host-specific at least to plant genus level, and that the inter-specific differences in male genitalia and color patterns of scutum, antenna, and maxillary palpus have contributed to reproductive isolation on the bryophytes that the flies share.

Shift of Feeding Mode in an Epizoic Stalked Barnacle Inducing Gall Formation of Host Sea Urchin.

Among diverse stalked barnacles, Rugilepas pearsei (Thoracica: Cirripedia: Arthropoda) is a rare unique species that is associated with echinoids and has highly atrophied cirri. We rediscovered the barnacle for the first time from description and verified that the barnacles live obligately in half-open galls formed on the test of the sea urchin Echinothrix diadema (Diadematidae: Echinodermata). A molecular phylogenetic analysis demonstrated that the obligate association with echinoids derived from epizoic life on crustaceans. A stable isotope analysis suggests that the barnacle feeds on particulate organic matter (POM) without parasitizing the host echinoids. These findings suggest that the host shift caused losses of plates and feather-like cirri, changes in the attachment device from cementation to anchoring, and a shift in feeding mode from filter feeding to POM collection. The barnacle's epizoic, superficially sub-endozoic, communal life in stout but narrow galls causes repetitive reproduction at the cost of reduced growth.

Morphological and ecological adaptation of limpet-shaped top shells (Gastropoda: Trochidae: Fossarininae) to wave-swept rock reef habitats.

Flattening of coiled shells has occurred in several gastropod lineages, while the evolutionary process of shell flattening is little known. The subfamily Fossarininae of the top shell family (Trochidae) is unique, because it includes four genera at various stages of shell flattening. Broderipia and Roya, have zygomorphic shells that has lost coiling, while the sister genera, Fossarina and Synaptocochlea, have respectively turbiniform and auriform shells. Therefore, comparisons of biology, habitats and detailed morphology among these four genera may help us to detect selection pressure driving shell flattening and loss of coiling. Although Broderipia has recently been identified as living symbiotically in the pits of sea urchins, the habitats and biology of the other three Fossarininae species, especially Roya are poorly known. After an extensive search on rocky shores of the Japanese Archipelago, we found live Roya eximia snails on intertidal/subtidal rock surfaces exposed to strong waves. Roya snails crept on the bare rock surface to graze periphyton at low tide, and fled into vacant barnacle shells at high tide. Comparison of the morphology of soft bodies in Fossarininae revealed that the columellar muscle of flattened species has been drastically elongated and arranged in posterior semi-outer edge of the flattened shell as observed in true limpets. The flattering and loss of coiling of the shell in Roya caused acquisition of a zygomorphic flat body, retraction of coiled visceral mass, and expansion of the foot sole. All of these changes improved tolerance against strong waves and the ability to cling to rock surfaces, and thus enabled a lifestyle utilizing both wave-swept rock surfaces and the inside of vacant barnacle shells.

A new large tellinid species of the genus Pharaonella from the Ryukyu Archipelago, Japan (Mollusca, Bivalvia).

A new tellinid species, Pharaonella amanyusp. n., is described from sand banks around Amami Islands, the Ryukyu Archipelago, in southern Japan. A molecular phylogenetic analysis suggests that this new species is closely related to P. sieboldii. This species has long siphons and lives buried deep in well-sorted white sand syntopically with Tonganaella tongana. These rare, large tellinid species are indicators of unspoiled tidal/subtidal sand flats, which should receive the highest priority conservation in the Ryukyu Archipelago.

Comparison of the Inducing Effect of Indole Compounds on Medusa Formation in Different Classes of Medusozoa.

Scyphozoa, Cubozoa and Hydrozoa are classes in the phylum Cnidaria that undergo metagenesis involving a dramatic morphological transition. In Scyphozoa and Cubozoa, when exposed to species- or strain-specific transition-inducing stimuli, asexually reproducing benthic polyps transform into sexually reproducing planktonic medusae. In Hydrozoa, exposure to species- or strainspecific transition-inducing stimuli causes formation of medusa buds in the polyp's body. In Aurelia aurita (Linnaeus, 1758) (Scyphozoa, Semaeostomeae), polyp-to-jellyfish transition is induced by some simple indole compounds. However, whether indole compounds can induce polyp-to-jellyfish transition in Cubozoa and Hydrozoa remains unknown. In the present study, we show that an indole compound, 5-methoxy-2-methylindole, induces polyp-to-jellyfish transition in Scyphozoa and Cubozoa. This inducing action suggests that the downstream steps of polyp-to-jellyfish transition are regulated by the same biochemical reactions in Scyphozoa and Cubozoa, irrespective of the type of transition-inducing environmental stimuli.