Cryptic species in Pacific sipunculans (Sipuncula: Phascolosomatidae): east‐west divergence between non‐sister taxa

Cryptic diversity, i.e. diversity observable in genetic but not in morphological traits, is prevalent in marine invertebrates and presents one of the greatest obstacles to obtaining accurate estimates of species richness. Sipunculans, commonly called peanut worms, are marine annelids in which high levels of cryptic diversity have previously been documented. In this study, we use genetic identification techniques to examine divergence of two lineages of Pacific sipunculans, both known under the name Phascolosoma agassizii. One lineage is isolated to the eastern Pacific coast while the other one inhabits the western Pacific coast. These clades are reciprocally monophyletic and are not recovered as sister taxa, suggesting relatively early divergence within Phascolosoma. Furthermore, we did not find support for a genetic distinction between the western Pacific Phascolosoma agassizii agassizii and Phascolosoma agassizii kurilense, a subspecies reported from the Kuril Islands. Considering that the type locality for P. agassizii is in the Eastern Pacific, we suggest that the western clade, including the samples from the Kuril islands, represent a new, undescribed species.     

Development of the tentacular apparatus in sipunculans (Sipuncula): I. Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920)

The development and arrangement of the tentacular apparatus of Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920) are described and illustrated using scanning electron microscopy. In T. nigra, the tentacular apparatus is composed of two crowns: the nuchal arc enclosing the nuchal organ and a crown of numerous oral tentacles arranged in U-shaped festoons. In early juveniles, two dorsal horn-like protrusions develop into the first, or primary, pair of tentacles of the nuchal arc. The second pair of tentacles of the nuchal arc develops dorsolaterally on the bases of the primary tentacles. Two ventrolateral lobes of the oral disk grow and become subdivided by the longitudinal ciliary groove into anlages of one set of dorsal and one set of ventral tentacles, thus forming a first oral festoon. Later, a pair of dorsolateral lobes develop between the first festoons and the nuchal arc to form a second pair of oral festoons. The third and following pairs of oral festoons develop in the dorsolateral growth zones lateral to the borders of the nuchal arc, where they meet the oral crown. The growing festoons extend down the oral disk and run alongside the head. A new oral tentacle appears directly at/on the base of the previous tentacle, thus giving rise to a typical sympodium with an alternating arrangement of tentacles. In T. pyroides, a second pair of tentacles develops from two ciliary lobes that are ventrolateral outgrowths of the circumoral ciliary field around the terminal mouth opening. The third pair of tentacles appears from the dorsolateral lobes at the base of primary tentacles, between the first two pairs of tentacles. These six tentacles determine the position of six main stems of the tentacular apparatus designated the first tentacles in the corresponding stems. The second tentacle in every stem appears as a ventrolateral outgrowth at the base of the first tentacle. The third and following tentacles in the stem are developed between the two previous tentacles according to a sympodial pattern. In both species, the distinct sympodial pattern in the arrangement of tentacles in the tentacular apparatus is well evidenced by the outlines of the ciliary oral grooves. The branched stems of T. pyroides may be homologized structurally and functionally to the oral festoons of T. nigra. J. Morphol. (c) 2006 Wiley-Liss, Inc.     

Deepsea herrings (Bathyclupeidae) of the northwestern Pacific ocean

Deepsea herrings of the northwestern Pacific (waters of Japan and Taiwan) are revised. Three species are recognized in this area, and two of them described as new: B. gracilis Fowler, 1938, B. japanotaiwana sp. n., and B. nikparini sp. n. B. japanotaiwana was previously mistakenly identified as the western Atlantic species B. argentea; actually it is especially similar to B. malayana, from which it differs in the bicolor oral cavity and in some meristic characters. B. nikparini is especially close to the Caribbean species B. schroederi, from which it differs in a greater number of developed gill rakers and smaller number of rays in the anal fin.     

Sipuncula of the southeastern Weddell Sea (Antarctica)

Sipunculans collected during the EPOS (European Polarstern Study) leg 3 cruise (13.1.89–10.3.89) to the Weddell Sea shelf area from depths ranging from 186 to 2037 m are recorded. Seven species in three genera are recognized; Nephasoma capilleforme and Phascolion convestitum have not been recorded hitherto in Antarctica. Significant morphological characters as shown by light (LM) and electron (SEM) photographs are described and illustrated. Antarctic distribution maps of the species collected and a checklist of the sipunculan species south of the Antarctic Convergence with depths and bibliographic references are included.     

Ultrastructure of the tentacles of Themiste lageniformis (Sipuncula)

Summary An ultrastructural study of the tentacles of Themiste lageniformis (Sipuncula) was conducted as part of a larger study of head metamorphosis in the species.The oral surface of the tentacles is constructed of a multiciliated, pseudostratified, columnar epithelium while the aboral surface is an unciliated, cuboidal epithelium. Intraepidermal mucous cells lie near the junction of the oral and aboral regions. The basal portion of the epidermal cells is embedded in a thick, collagenous extracellular matrix which contains outer circular muscles, inner longitudinal muscles, the main tentacular nerve and its branches. Three tentacular canals are present and are lined by peritoneum. Hemerythrocytes and coelomocytes flow through the lumen of the canals in a regular pattern.Justification for the designation of the tentacular canals as coelomic rather than vascular is discussed.     

Phylogeography of the marine macroalga Sargassum hemiphyllum (Phaeophyceae,Heterokontophyta) in northwestern Pacific

Sargassum hemiphyllum is commonly found in Japan and Korea, with a variety, var. chinense, that is found distributed in the southern Chinese coast. We previously reported distinct genetic differentiation between the two taxa based on the PCR‐RFLP data of plastid RubiscoL‐S spacer. The present study aims at elucidating the phylogeographic pattern of S. hemiphyllum based on more markers in the nuclear and extranuclear genomes, with a view to reveal the occurrence of hybridization. The two allopatrically distributed taxa were found to be genetically distinct in nuclear ITS2, plastidial Rubisco (Rbc) and mitochondrial TrnW_I (Trn) spacers. Their divergence was postulated to be attributable to the vicariant event which resulted from the isolation of the Sea of Japan during the late Miocene (6.58–11.25 Mya). Divergence within both S. hemiphyllum and the chinense variety was observed based on Trn spacer, while the divergence in S. hemiphyllum was further confirmed in Rbc spacer. This divergence appears to correspond to the separation of the Japanese populations between the Sea of Japan and the Pacific that occurred around 0.92–2.88 Mya (the early Pleistocene). The presence of an ITS2 clone resembling var. chinense sequences in a Japanese population of S. hemiphyllum (JpNS) raises the possibility of the introgression of var. chinense individuals into S. hemiphyllum population. Compared to that between S. hemiphyllum and the chinense variety, hybridization among the Japanese and Korean populations of S. hemiphyllum is highly probable as all these individuals share a pool of nuclear ITS2 sequences, possibly attributable to incomplete concerted evolution of ITS2.     

A revision of the genera Sipunculus and Xenosiphon (Sipuncula)

A critical analysis of historically used characters, based on a large series of specimens, is presented. Based on a comparative morphological analysis of type specimens, the 19 species of Sipunculus and four of Xenosiphon are reviewed. The genera and subgenera are redefined so that, after appropriate revisions arc made, Sipunculus (Sipunculus) has eight, Sipunculus (Contraporus) new subgenus has two, and Xenosiphon has only one remaining valid species. A zoogeographical note and a key to the species are included.     

Abyssal benthic foraminifera from the northwestern Pacific (Shatsky Rise) during the last 298 kyr

Benthic foraminifera in a gravity core from Shatsky Rise (northwestern Pacific, water depth 2612 m) show large fluctuations in accumulation rate, species composition and diversity over the last 298 kyr. The most important fluctuations (explaining more than 90% of the faunal variance) result from variations in relative abundance of the three most abundant species: Epistominella exigua, Alabaminella weddellensis and Uvigerina peregrina. High accumulation rates of U. peregrina, a species linked to high, continuous productivity, occurred with a 100 kyr periodicity at the end of glacial stages, and correspond to high mass accumulation rates of organic carbon. Peak accumulation rates of E. exigua occurred during glacial stage 4 and the middle part of glacial stages 8 and 6, whereas A. weddellensis was dominant in the early part of stages 8 and 6, and the late part of stage 3. A high abundance of these species probably indicates lower overall productivity than a high abundance of U. peregrina, and their relative and absolute abundances may be linked not simply to the amount of organic matter delivered to the sea floor, but to the intermittent delivery of fresh, easily degraded organic matter. The overall species diversity is negatively correlated to the relative abundance of E. exigua, but not to that of A. weddellensis, implying that these species have different environmental preferences, although both have been linked to periodic phytodetritus deposition in the present oceans.     

Larvae of Pterothrissus gissu (Albilidae) in temperate waters of the northwestern part of the Pacific Ocean

According to results of trawl and oceanological surveys, the distribution and numbers of leptocephalic larvae of Japanese gissu Pterothrissus gissu in June–July 2010–2011 in temperate waters of the northwestern part of the Pacific Ocean in connection with water circulation and distribution of thermohaline characteristics are considered. Larvae of Japanese gissu in the water area of the survey were characterized by high occurrence and numbers; all of them were caught in a layer of 0–30 m mainly in the dark hours of the day at the temperature of the surface water layer of 2.8–8.7°C. Their distribution in waters of the western subarctic circulation is related to both constant and wind currents.     

Photoreceptive organs in larvae of Spionidae (Annelida) and Sipuncula

A pair of spherical unpigmented ocelli in addition to pigmented eyes have been observed in the anterior part of the prostomium in larvae of more than 40 species of Spionidae examined alive with light microscopy. Ocelli become visible in larvae at the one- to three-segment stage, increase in size as growth proceeds, and probably disintegrate in the course of metamorphosis because they were not observed in adults and settled juveniles. One pair of transparent, spherical bodies is also found in the anterior part of the head of planktotrophic pelagosphera larvae of Sipuncula. These bodies are in a similar position and have a similar appearance and size to unpigmented ocelli in Spionidae larvae. A pair of epidermal invaginations, densely covered with short cilia, is also observed antero-laterally in the head in pelagospheras of one species. These invaginations appear similar to the nuchal organs present in many polychaete larvae. Photoreceptive organs so far reported for sipunculan larvae comprise only pigmented eyes. Unpigmented ocelli and nuchal organs have never been reported in pelagospheras. Further ultrastructural investigations on sipunculan larvae are encouraged to clarify the composition, function and morphogenesis of transparent, spherical bodies and ciliated invaginations in the anterior part of the head. Such investigations may help to better understand the nature of photoreceptive structures and nuchal organs in Sipuncula, and also contribute to phylogenetic hypotheses regarding relationships of the Sipuncula and Annelida.     

Biological control of the wetlands weed purple loosestrife (Lythrum salicaria) in the Pacific northwestern United States

Purple loosestrife (Lythrum salicaria) is an Eurasian perennial hydrophyte that has become naturalized in wetlands and in and along waterways throughout temperate North America. The ecological integrity of such areas is threatened by rapidly forming monotypic infestations that displace valued flora and diminish critical fish and wildlife habitat. The inability of physical, cultural, and chemical methods to provide adequate control of the weed has led to the development of an insect-based biological control program. The first field releases of the bud and leaf feeding beetles, Galerucella calmariensis and G. pusilla, and a root-mining weevil, Hylobius transversovittatus, were made in the United States and Canada in 1992. A total of 4740 Galerucella spp. adults were released in central Washington during 1992 and 1993 at eight sites and 471 H. transversovittatus egg inoculations were made in 1993 at three locations. Establishment of both Galerucella spp. was confirmed and Hylobius colonization was achieved.     

Comparative phylogeography in marginal seas of the northwestern Pacific

The maturation of marine phylogeography depends on integration of comparative information across different regions globally. The northwestern Pacific, characterized by unique tectonic setting, however, is still underrepresented. This study seeks to highlight its phylogeographical history based on the available population data, focusing on three seas: the East China Sea (ECS), the South China Sea (SCS) and the Sea of Japan (SOJ). We first conducted a literature survey to evaluate current research efforts and then reanalysed the population structure, historical demography and genealogy for two selections of studies (namely ‘the ECS category’ and ‘the multiple‐sea category’) to elucidate the evolutionary processes within and across the seas, respectively. For the ECS category, the meta‐analyses revealed most studies displayed a shallow phylogeny, indicating a single origin from the sea. Significant population structure was commonplace, particularly in molluck and crustacean studies, with proportions of 89% and 80%, respectively. Nearly all studies selected showed signals of population expansion: the times estimated were closely linked to a period of ~120–140 Kya rather than the last glacial maximum. For the latter category, divergent intraspecific lineages appeared among seas and overlapped in the adjacent regions, a pattern implying each sea had served as an independent refugium during glaciations. The genetic splits, however, were estimated to arise from separate events dating from late Miocene to middle Pleistocene. As phylogeography is still in its infancy in the region, more effort is needed to test and complement the general rules abstracted here. Finally, challenges and prospects were discussed to accelerate further research.     

Genetic structure in the marbled rockfish (Sebastiscus marmoratus) across most of the distribution in the northwestern Pacific

The marbled rockfish (Sebastiscus marmoratus) is an important species that is widely distributed across the marginal seas of the northwestern Pacific. Several kinds of DNA markers, such as amplified fragment length polymorphisms (AFLP), mitochondrial DNA (mtDNA) and single nucleotide polymorphisms (SNPs), have been used to assess the population genetic characteristics of this species in previous studies. However, there have been no genetic profiling studies to cover the entire distribution. Here, six highly polymorphic microsatellite markers were applied to 18 populations across a spatially large area (from the Sea of Japan in the north to the South China Sea in the south). The results showed that the whole population exhibits high, stable genetic diversity, low relatedness and large effective population size. There was very weak genetic differentiation overall and no isolation by distance occurred among the populations, which may be attributed to significant contemporary gene flow, each generation. The above results may lead us to infer that the recent S. marmoratus population structure could be considered as one large integrated population. The present study will be beneficial to population conservation and fisheries management of S. marmoratus and will lead to better insights into the genetic characteristics of other species that belong to the Sebastidae family.     

Saurida fortis,a new species of lizardfish (Aulopiformes: Synodontidae) from the northwestern Pacific Ocean

Ichthyological Research - Saurida fortis sp. nov., a lizardfish (Aulopiformes: Synodontidae) from the northwestern Pacific Ocean, previously misidentified as Saurida umeyoshii Inoue and Nakabo 2006...     

Fine structure of the nephridial muscle layer cells of Phascolosoma granulatum (Sipuncula)

The nephridial muscle layer of Phascolosoma granulatum consists of a network of longitudinal and circular cells separated by connective tissue matrix. The muscle fibers are densely packed with thick and thin myofilaments, among which are scattered cytoplasmic dense bodies. The nucleus and noncontractile cytoplasmic organelles occupy a lateral projection from the contractile portion of the fiber. Cytoplasmic dense bodies are the result of a clustering of an indeterminate number of the thin actin filaments that fill the cytoplasm between thick filaments. Attached to the cytoplasmic face of the cell membrane are membrane-associated electron-dense plaques. These sites are linked to the contractile myofilaments by narrow filamentous bridges. Extracellular narrow filaments extend from these plaques to collagen fibers of the connective tissue matrix. Differences in length of the dense plaques may be related to differences in thick myofilament diameter in three types of muscle fiber, types A, B and C, statistically distinguished by mean fiber size differences. The plaques may serve as connecting links for the transmission of tension from contractile units to the connective tissue of the muscle layer. © 1993 Wiley-Liss, Inc.     

An examination of the Phascolosotna subgenera Antillesoma, Rueppellisoma and Satonus (Sipuncula)

Three of the subgenera in the sipunculan genus Phascolosoma are reviewed, based on examination of type material. The presumed difference between the subgenera Antillesoma and Rueppellisoma (number of retractor muscles) is shown to be invalid and the taxa are therefore synonymous. The 15 species are either considered junior synonyms of P. antillarum (eight), transferred to other species or genera (six), or considered incertae sedis (one). Phascolosoma antillarum is redescribed. Of the eight species in the subgenus Satonus, only P. pectinatum remains valid: the others are considered either to belong to other taxa (four) or to be incertae sedis (three).     

The characteristics of feeding of Diaphus theta (Myctophidae) in the northwestern part of the Pacific Ocean in the Summer-Autumn period

The feeding of Diaphus theta in the zone of the subarctic front and in Pacific waters off the Kuril Islands in November 2001 and in July–August 2002 was studied. Interzonal plankton [euphausiids (mainly Euphausia pacifica), copepods (Metridia pacifica), and amphipods (Themisto pacifica)] dominated in the food bolus. As fish grow, the proportion of copepods decreases, the proportion of euphausiids and amphipods increases in stomachs, and the food spectrum extends. One considerable peak in the feeding (in both the summer and autumn periods) occurring in the nighttime was revealed. The most intensive feeding takes place in the dark time of the day in the epipelagial due to euphausiids and amphipods; in the light time of the day, the feeding activity declines. Fish feed mainly on copepods in the upper pelagial. The daily ration of D. theta was 1.5% of the weight of fish in the summer period and 3.2% in the autumn period.     

Reconstructing the phylogeny of the Sipuncula

Sipunculans are marine spiralian worms with possible close affinities to the Mollusca or Annelida. Currently 147 species, 17 genera, 6 families, 4 orders and 2 classes are recognized. In this paper we review sipunculan morphology, anatomy, paleontological data and historical affiliations. We have conducted cladistic analyses for two data sets to elucidate the phylogenetic relationships among sipunculan species. We first analyzed the relationships among the 45 species of Phascolosomatidea with representatives of the Sipunculidea as outgroups, using 35 morphological characters. The resulting consensus tree has low resolution and branch support is low for most branches. The second analysis was based on DNA sequence data from two nuclear ribosomal genes (18S rRNA and 28S rRNA) and one nuclear protein-coding gene, histone H3. Outgroups were chosen among representative spiralians. In a third analysis, the molecular data were combined with the morphological data. Data were analyzed using parsimony as the optimality criterion and branch support evaluated with jackknifing and Bremer support values. Branch support for outgroup relationships is low but the monophyly of the Sipuncula is well supported. Within Sipuncula, the monophyly of the two major groups, Phascolosomatidea and Sipunculidea is not confirmed. Of the currently recognized families, only Themistidae appears monophyletic. The Aspidosiphonidae, Phascolosomatidae and Golfingiidae would be monophyletic with some adjustments in their definition. The Sipunculidae is clearly polyphyletic, with Sipunculus nudus as the sister group to the remaining Sipuncula, Siphonosoma cumanense the sister group to a clade containing Siphonosoma vastumand the Phascolosomatidea, and Phascolopsis gouldi grouping within the Golfingiiformes, as suggested previously by some authors. Of the genera with multiple representatives, only Phascolosoma and Themiste are monophyletic as currently defined. We are aiming to expand our current dataset with more species in our molecular database and more detailed morphological studies.