Authorship：Corresponding author   Publishing type：Research paper (scientific journal)  

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Authorship：Lead author,　Corresponding author   Language：English   Publishing type：Research paper (scientific journal)  

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：CAMBRIDGE UNIV PRESS  

Nymphonella tapetis (Pycnogonida, Ascorhynchidae) is an endoparasitic sea spider affecting bivalves. Recently, sea spiders have been found on a massive scale in the commercially important Manila clams (Veneridae, Ruditapes philippinarum) in Japan (Tokyo Bay). Simultaneously, mass mortality has occurred in this area. Local fishers assumed that this mass mortality was caused by the parasitic sea spider, despite the effect of the parasite and parasite intensity on the host being unknown. To evaluate the susceptibility of the Manila clam to sea spider infestation and the impact on mortality levels, we established six treatments at different infection intensities (density of newly hatched larvae of sea spiders) over a 6-month long laboratory experiment. We monitored mortality and three susceptibility indices (clearance rate, sand-burrowing speed and adductor muscle strength) under sufficient food conditions. Parasitization by sea spider affected clearance rate and sand-burrowing speed. The pattern of parasitic intensity effects on survival of Manila clam hosts was shown to be dependent on the levels of parasite numbers, i.e. clams with lower parasitic levels (total of <200 hatching larvae of sea spider given to a host) have a higher survival rate, and high mortality of host clams was shown in excessively higher parasitic densities (400-4000 individuals). Such pattern of parasitic effects on host survival might be one of the causes of mass mortality of Manila clams occurring in the field.

DOI： 10.1017/S0025315417000200

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：TAYLOR & FRANCIS LTD  

Postembryonic developmental stages of an endoparasitic pycnogonid, Ammothella biunguiculata in Izu Peninsula, Japan are described. Eleven stages were identified beginning with a protonymphon larva attached to the male oviger. We found endoparasitic individuals in the host actinian from the second to tenth instar, and forms in the ninth stage to adult were found free-living. This indicates a transition from being endoparasitic to free-living during the ninth to tenth instar stages. The first instar protonymphon attached to the adult male oviger has a gland duct on the anterior margin of each chelifore scape which completely disappears with the second instar. The disappearance of the chelifore gland duct coincides with the beginning of an endoparasitic stage in the development of this species. Although the larval morphology and the postembryonic development of pycnogonids have been summarized by several authors, the present study concludes that much remains to be learnt.

DOI： 10.1080/07924259.2017.1291452

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：SPRINGER  

A sudden outbreak of the parasitic sea spider Nymphonella tapetis accompanied by mass mortality of the Manila (asari) clam Ruditapes philippinarum was observed in Japan in 2007. It was unknown why dead and dying clams were found on the sand surface. In our laboratory experiment, in which sea spiders were artificially parasitised to host clams, unburied clams were significantly increased, suggesting that sea spider parasitism can change the burial mode of clams. We compared parasite loads, somatic condition and reburial activity of Manila clams among three burial groups (fully buried, partly buried and unburied) collected from the field. Unburied and partly buried clams had 30% lower soft body weight and 60% lower adductor strength as compared to the fully buried clams. Parasite prevalence and the mean infection intensity were greater in unburied and partly buried clams than in fully buried clams. Reburial depth was smaller in unburied and partly buried clams than in fully buried clams under laboratory conditions. Approximately 30% of the unburied and partly buried clams died during 30 days, whereas fully buried clams suffered no mortality. These results indicate that sea spider parasitism can reduce the burial depth and somatic condition of host clams, and eventually cause clam mortality on the sand.

DOI： 10.1007/s10750-015-2564-y

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：Seto Marine Biological Laboratory, Kyoto University  

DOI： 10.5134/217458

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：CRUSTACEAN SOC  

We obtained the nucleotide sequence of 18S rDNA of Nymphonella tapetis, a pycnogonid endoparasitic on some bivalves, together with that of other species belonging to Ascorhynchidae Hoek, 1881. The phylogenetic tree based on these sequences as well as those of selected species from a database strongly supports the monophyly of Ascorhynchidae and the inclusion of Nymphonella in the family. Nymphonella seems to be an aberrant form in the Ascorhynchidae, but multi-gene analysis with more species is required for the exact determination of the taxonomic position of the genus.

DOI： 10.1163/1937240X-00002348

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：TAYLOR & FRANCIS LTD  

Reproductive traits of scleractinian corals are important in understanding their ecology and for maintaining coral populations; however, it has been reported for no more than 30% of species. Oulastrea crispata (Lamark, 1816) has the greatest north-south distribution in the Indo-West Pacific. This species is also known as one of the most stress-tolerant corals. Thus, most previous studies on O. crispata have examined its physiological, ecological, morphological, or genetic traits and its symbiotic dinoflagellates relative to its broad distribution and its stress tolerance. However, there have been few studies on its reproduction. In this study, the release of sperm and eggs of O. crispata was observed under laboratory rearing conditions in Shirahama, Kii Peninsula, Japan. This is the first direct evidence of sexual reproduction in this species, and the contribution of sexual reproduction to Oulastrea biology is briefly discussed.

DOI： 10.1080/07924259.2015.1006340

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：ACAD SINICA INST ZOOLOGY  

Background: Goniopora stokesi is an uncommon species mainly found in tropical and subtropical regions but was also reported in temperate regions of Japan. This species has two unique characteristics. First, it does not typically attach to hard substrates, i.e., it is free-living. Although non-free-living colonies were reported, it is unclear whether their morphologies exhibit intraspecific or interspecific differences. Second, they can asexually form daughter colonies, which are secondary colonies that grow on the surface (coenosac) of a parent colony and subsequently detach. To date, this specific characteristic has not been reported in Japan, and it is not clear whether this species occurs in Japan.
Results: To clarify the taxonomic status of this species, we investigated its reproductive mechanism in Japan and morphologically and genetically analyzed specimens collected from both subtropical and temperate regions. We found that this species forms daughter colonies in Okinawa, Japan and that free-living colonies in the temperate region, which were formerly recognized as G. stokesi, likely constitute a morphological variation of a separate species. In addition, all non-free-living colonies with G. stokesi-like morphologies were also morphological variations of other species.
Conclusions: Overall, free-living colonies with large, deep calices, thin walls, and well-developed epitheca on the underside were G. stokesi, whereas other free-living and all non-free-living G. stokesi-like colonies belonged to other species, such as Goniopora djiboutiensis.

DOI： 10.1186/1810-522X-52-25

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：CAMBRIDGE UNIV PRESS  

A new marine tardigrade of the genus Tanarctus is described from Japan. Tanarctus diplocerus sp. nov. was collected from Shimabara Bay, Kumamoto and Tanabe Bay, Wakayama. It is characterized by simple leg IV appendages, the lack of the internal-external relation in the dorsal and ventral cirri and the lack of the club-shaped secondary clavae. The secondary sexual dimorphism of the primary clavae and probable two-digit juveniles of the new species are reported. The problematic terminology of the so-called 'internal' and 'external' cirri is also discussed. A taxonomic key for the species of Tanarctus is provided.

DOI： 10.1017/S0025315412000689

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：ZOOLOGICAL SOC JAPAN  

A new species of the previously monospecific marine tardigrade family Neostygarctidae is described. Neostygarctus lovedeluxe n. sp. was found from a submarine cave in Miyako Islands, Japan. This is the first record of Neostygarctidae from the Pacific. The new species is easily distinguished from the previously known N. acanthophorus by its number of dorsal spines, as N. lovedeluxe has two spines each on the three dorsal body plates in contrast to one in N. acanthophorus. Furthermore the morphology of the two clawed juvenile is reported for the first time in Neostygarctidae, providing new insights into the common sequence in some ontogenic traits.

DOI： 10.2108/zsj.30.414

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：Academia Sinica  

Background: Goniopora stokesi is an uncommon species mainly found in tropical and subtropical regions but was also reported in temperate regions of Japan. This species has two unique characteristics. First, it does not typically attach to hard substrates, i.e., it is free-living. Although non-free-living colonies were reported, it is unclear whether their morphologies exhibit intraspecific or interspecific differences. Second, they can asexually form daughter colonies, which are secondary colonies that grow on the surface (coenosac) of a parent colony and subsequently detach. To date, this specific characteristic has not been reported in Japan, and it is not clear whether this species occurs in Japan. Results: To clarify the taxonomic status of this species, we investigated its reproductive mechanism in Japan and morphologically and genetically analyzed specimens collected from both subtropical and temperate regions. We found that this species forms daughter colonies in Okinawa, Japan and that free-living colonies in the temperate region, which were formerly recognized as G. stokesi, likely constitute a morphological variation of a separate species. In addition, all non-free-living colonies with G. stokesi-like morphologies were also morphological variations of other species. Conclusions: Overall, free-living colonies with large, deep calices, thin walls, and well-developed epitheca on the underside were G. stokesi, whereas other free-living and all non-free-living G. stokesi-like colonies belonged to other species, such as Goniopora djiboutiensis. © 2013 Kitano et al.
licensee Springer.

DOI： 10.1186/1810-522X-52-25

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：INT SCIENCE SERVICES/BALABAN PUBLISHERS  

The postembryonic development of the female reproductive system in the pycnogonid Propallene longiceps is examined. The germ cells can be detected first in the later stage of the 3rd instar and become a paired gonad covered with gonadal epithelium in the next instar. The larval gonad changes its shape: paired at the 4th instar, reversed U-shaped at the fifth, unpaired at the sixth, and paired again at the seventh. Oocytes can be distinguished, and the extension of the ovary into the walking legs begins at the 7th instar. Growing oocytes protrude outward from the ovary on cellular stalks in the pedal part. The trunk ovary becomes U-shaped, and the oviducts and genital pores start forming at the 8th instar. The disappearance of trunk ovary begins at the 9th instar, and is complete at the next adult stage. The connection between the pedal ovarian lumen and the genital pores via the oviducts is complete in the adult, and the female reproductive system becomes segmentally arranged. This study confirms that the segmental arrangement of adult female reproductive system in P. longiceps, which is unique among recently described arthropods, is a secondary state in pycnogonids attained by reducing the trunk part of ovary.

DOI： 10.1080/07924259.2011.613167

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：MARINE BIOLOGICAL LABORATORY  

In the lepadomorph barnacle Octolasmis warwickii, individuals are often found attached to the scutum of conspecifics living externally on the crab hosts. To test whether these conspecific-attached individuals are dwarf males, as are known to occur in other suborders of barnacles, we compared the pattern of attachment, size-frequency distribution, and reproductive status of the conspecific-attached (Con-A) and crab-attached (Crab-A) individuals. Con-As were smaller than Crab-As. There was a positive relationship between the body size of Crab-As and the number of individuals on them. Con-As had longer penises than Crab-As of the same body size, and their testes were better developed. The four largest Con-As examined were brooding eggs. These results indicate that Con-As of O. warwickii are dwarf males, with a potential to become hermaphroditic. This represents the first known example of coexistence of males and hermaphrodites in the suborder Lepadomorpha. The mating group size of O. warwickii was smaller than in its hermaphroditic congeners but larger than in barnacles with dwarf males and females, which supports the current theories that group size is important for the evolution of sexuality patterns in barnacles.

DOI： 10.1086/BBLv218n3p259

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The structure of the adult female reproductive system in the pycnogonid. Propallene longiceps, is examined. The ovary is present in each walking leg from the second coxa to femur, containing oocytes in various developmental stages. Oocytes are protruded outward from the ovary on cellular stalks. Each ovary is directly connected with the genital pore through the oviduct in the second coxa. Thus ovulation of the mature ova is into the ovarian lumen and oviposition is through the oviduct and the genital pore. Such a route of ovulation and oviposition has been commonly described in other pycnogonids as well as in euchelicerates. However, the segmental arrangement of the female reproductive system revealed in the present study, with isolation of a complete set of ovary, oviduct and genital pore in each walking leg, is distinct from the situation described in other pycnogonids, and recalls a hypothesized ancestral state in arthropods.

DOI： 10.1080/07924259.2010.9652314

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：ZOOLOGICAL SOC JAPAN  

The phylum Mollusca is one of the major groups of Lophotrochozoa. Although mollusks exhibit great morphological diversity, only a few comparative embryological studies have been performed on this group. In the present study, to begin understanding the molecular development of the diverse morphology among mollusks, we observed early embryogenesis in a bivalve, the Japanese spiny oyster, Saccostrea kegaki. Although several studies have begun to reveal the genetic machinery for early development in gastropods, very little molecular information is available on bivalve embryogenesis. Thus, as a step toward identifying tissue-specific gene markers, we sequenced about 100 cDNA clones picked randomly from a gastrula-stage cDNA library. This basic information on bivalve embryology will be useful for further studies on the development and evolution of mollusks.

DOI： 10.2108/zsj.25.455

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Language：English   Publishing type：Research paper (scientific journal)   Publisher：ELSEVIER SCI LTD  

Vitellogenic oocytes of all pycnogonids studied so far contain dilated elements of the endoplasmic reticulum, filled with characteristic electron-dense bodies. During vitellogenesis these bodies fuse and form larger, almost spherical granules that were traditionally interpreted as nascent yolk granules. Here, we present the results of ultrastructural investigations of previtellogenic and early vitellogenic oocytes of Propallene longiceps (Pycnogonida, Callipallenidae). We show that the intra-cisternal bodies/granules of pycnogonids are not involved in vitellogenesis but Contain macromolecules that are released from the oocyte and contribute to the formation of an egg envelope. The obtained results are discussed in a phylogenetic context. We suggest that the presence of the intra-cisternal electron-dense bodies in the oocyte cytoplasm represents a plesiomorphic character of arthropods inherited from the arthropod ancestor. (C) 2007 Elsevier Ltd. All rights reserved.

DOI： 10.1016/j.asd.2007.06.001

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Ovarian ultrastructure and oogenesis in two pycnogonid species, Cilunculus armatus and Ammothella blunguiculata, were investigated. The ovary is morphologically and functionally divided into trunk and pedal parts. The former represents the germarium and contains very young germ cells in a pachytene or postpachytene phase, whereas the latter houses developing previtellogenic and vitellogenic oocytes and represents the vitellarium. Intercellular bridges were occasionally found between young (trunk) germ cells. This indicates that in pycnogonids, as in other animal groups, at the onset of oogenesis clusters of germ cells are generated. As nurse cells are absent in the ovaries of investigated species, the clusters must secondarily split into individual oocytes. In the vitellarium, the oocytes are located outside the ovary. Each oocyte is connected to the ovarian tissue by a stalk composed of several somatic cells. The stalk cells directly associated with the oocyte are equipped with irregular projections that reach the oocyte plasma membrane. This observation suggests that the stalk cells may play a nutritive role. The ooplasm of vitellogenic oocytes comprises mitochondria, free ribosomes, stacks of annulate lamellae, active Golgi complexes, and vesicles derived from these complexes. Within the latter, numerous electron-dense bodies are present. We suggest that these bodies contribute to yolk formation. © 2006, The American Microscopical Society, Inc.

DOI： 10.1111/j.1744-7410.2006.00066.x

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The ascidian nervous system is extremely simple, although the structure of it is comparable with the complex vertebrate nervous system. This simplicity makes the ascidian nervous system a good model to understand how the neuronal circuit is built up in the chordate nervous system. In order to study the formation of the neuronal circuit at the single cell level, molecular markers to characterize specific single cells are desired. In the present paper, we describe the gene expression pattern of CiGl: an ascidian homologue of Goliath, a Drosophila RING-finger gene. In the early embryonic stage, CiGl is expressed in the lateral part of the neural tube and in several peripheral nerve cells. Later in the larval stage, CiGl specifically marks ocellus: one of the pigment cells in the ascidian brain, which is involved in the photoreceptive system. CiGl will be useful to understand the differentiation mechanism of ocellus, and especially to test the model proposed by Darras and Nishida (2001). In addition, the finding of this single cells specific gene expression pattern at a certain developmental stage encourages us to look for more genes which mark single cells, especially those that have not been well characterized. (C) 2003 Elsevier Science B.V. All rights reserved.

DOI： 10.1016/S0378-1119(03)00606-1

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The shape of foregut lumen in both the pharyngeal and the oesophageal regions has been examined from serial histological sections in 16 species of pycnogonids covering all extant families. Each species has a Y-shaped foregut lumen in transverse section. This result shows that some previous authors misoriented the lumen as inverted Y-shaped. The triradiate Y-shaped foregut lumen has been found among ecdysozoans including tardigrades, onychophorans and scatteringly arthropods. At present, it is not clear whether the Y-shaped state of the characteristic in pycnogonids is convergent or symplesiomorphy with that in tardigrades and onychophorans.

DOI： 10.1017/S0025315402006628

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The structure of the adult female reproductive system was examined in a schizomid, Schizomus sawadai. The system consists of a single median ovary, a pair of lateral oviducts, and 4 single common oviduct with a pair of triplets of small seminal receptacles and a simple slit-like genital pore. Smaller previtellogenic oocytes less than 40 gm in diameter are found sparsely scattered in the dorsal ovarian wall among the ovarian epithelial cells, while two size groups of advanced oocytes, larger previtellogenic and vitellogenic, protrude from the lateral and ventral ovarian walls, respectively, accompanied by short cellular stalks. Neither nurse cells nor follicle cells are present around all the oocytes. Oogonia are not found in the adult ovary, indicating the lack of a continued supply of oocytes. The ovarian lumen is connected with the lumens of the lateral and common oviducts, and then with the genital pore. Modes of oogenesis, ovulation, and oviposition, deduced from the structural features of the female reproductive system of S. sawadai, are compared with those in some other arachnids, merostomes and pycnogonids.

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Transgenic fish bearing the mouse tyrosinase gene (mg-Tyrs-J) were produced by transfection into fertilized eggs of the homozygous normal orange-colored variant of medaka fish, Oryzias latipes, by means of electroporation. Of 589 eggs transfected, 38 fish (6%) exhibited brownish wild-type skin pigmentation, which was discernible from control siblings. Light microscopy of the skin from the founders thus generated disclosed that 1) melanization occurred and was restricted to melanophores formed presumably from preexisting amelanotic melanophores, 2) there was a wide variation in the degree of melanization observed among melanophores, and 3) no melanin deposition was recognized in xanthophores or leucophores. Immunofluorescence using an antibody raised against mouse tyrosinase disclosed that melanophores at varying stages of maturation were reactive. Thus, it was shown that the transgene in medaka fish expressed its action in a cell type-specific manner. Crossing of transgenic founders with homozygous orange-colored variant fish yielded two groups of offspring expressing either the wild-type or the orange-colored skin pigmentation at an approximate ratio of 1:1. Crossing between founders exhibiting wild-type pigmentation yielded only offspring with melanized skin. Skin melanophores in these offspring formed vertical stripes, which are rare in this species. The hereditary basis of melanized skin was demonstrated in matings of F1 progenies, which resulted in similar degrees of melanization over whole skin melanophores. The sum of these findings implied that the transgene is expressed as a dominant character gene and is transmitted through germ cell lines according to the Mendelian law. PCR analysis combined with nested PCR technique strongly suggested that the transgene was integrated into the medaka genome, even though the copy number deduced from gel banding was largely diminished, possibly as a result of fragmentation or instability within the medaka genome.

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DOI： 10.12782/specdiv.1.117

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DOI： 10.5134/176249

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An intersexual specimen of the sea spider, Cilunculus armatus, having female and male characteristics together with some intermediate ones was collected from Sagami Bay. Eggs are seen through the cuticle of each femoral segment of walking legs. At the same time, cement gland, peculiar to male, is also found on the same segment of each walking leg, but somewhat atrophied and distally shifted. Genital pore opening on the second coxal segment of each walking leg as seen in female is of intermediate size between the sizes of both sexes. Oviger is intermediate in size and shape, and the number and distributional pattern of ovigerous compound spines are also intermediate.

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As in other pycnogonids, the adult female reproductive system of Endeis nodosa is limited to the trunk and walking legs of the cephalothorax. The U‐shaped trunk ovary extends eight blind branches separately into the corresponding walking legs up to each femoral segment. In both the trunk and pedal region, the ovary lies dorsally above the gut, sandwiched between the dorsal and the ventral layer of the horizontal septum. Oogonia and very young oocytes occur throughout the ovary. They fill the trunk ovary and, in the tubular pedal ovary, are aggregated into several cord‐shaped longitudinally arranged germ zones. More advanced oocytes occur only in the pedal ovary. None lie in the germ zones. but instead they protrude individually from the wall of the pedal ovary into the hemocoel on cellular stalks. Vitellogenesis occurs in the larger stalked oocytes. An oval genital pore with a cuticular lid lies on the ventral surface of the second coxa of each walking leg. The pedal ovarian lumen is directly connected to the genital pore by a short oviduct in the second coxa. The ovarian structure and oogenetic modes in this pycnogonid species show some common characteristics with those of most chelicerates. Copyright © 1991 Wiley‐Liss, Inc.

DOI： 10.1002/jmor.1052090303

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