Long-term occurrence of hybrids between Japan Sea and Pacific Ocean forms of threespine stickleback, <Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>, in Hokkaido Island,Japan

The distribution patterns of two genetically divergent forms (Japan Sea and Pacific Ocean forms) of Gasterosteus aculeatus and their hybrids in the Hokkaido Island, Japan were investigated. Hybrid frequencies and the mating pattern of natural F1-hybrids and backcrosses, were examined by using allozyme analysis. The distribution patterns matched those previously reported. All three sympatric localities in eastern Hokkaido Island included hybrids between the two forms in each year examined. A low frequency of hybrids of sticklebacks has been maintained in areas of sympatry of the two forms in Hokkaido Island for several years. The proportion of F1-hybrids was significantly lower than expected by random mating in six samples from four sympatric populations, suggesting form-assortative mating. Mating crosses of natural F1-hybrids existed in both patterns. Natural F1-hybrids backcrossed more frequently with Pacific Ocean form than Japan Sea form. However, the proportions of backcross individuals in two of three samples were not significantly different from the expected proportions by random mating between F1-hybrids and their parent forms. In other samples, despite the Pacific Ocean form predominant in a population, F1-hybrid predominantly backcrossed with the Japan Sea form. These results support the hypothesis of gene flow from Pacific Ocean to Japan Sea forms.     

Comparative Analysis of Japanese Three-Spined Stickleback Clades Reveals the Pacific Ocean Lineage Has Adapted to Freshwater Environments while the Japan Sea Has Not

Divergent selection and adaptive divergence can increase phenotypic diversification amongst populations and lineages. Yet adaptive divergence between different environments, habitats or niches does not occur in all lineages. For example, the colonization of freshwater environments by ancestral marine species has triggered adaptive radiation and phenotypic diversification in some taxa but not in others. Studying closely related lineages differing in their ability to diversify is an excellent means of understanding the factors promoting and constraining adaptive evolution. A well-known example of the evolution of increased phenotypic diversification following freshwater colonization is the three-spined stickleback. Two closely related stickleback lineages, the Pacific Ocean and the Japan Sea occur in Japan. However, Japanese freshwater stickleback populations are derived from the Pacific Ocean lineage only, suggesting the Japan Sea lineage is unable to colonize freshwater. Using stable isotope data and trophic morphology, we first show higher rates of phenotypic and ecological diversification between marine and freshwater populations within the Pacific Ocean lineage, confirming adaptive divergence has occurred between the two lineages and within the Pacific Ocean lineage but not in the Japan Sea lineage. We further identified consistent divergence in diet and foraging behaviour between marine forms from each lineage, confirming Pacific Ocean marine sticklebacks, from which all Japanese freshwater populations are derived, are better adapted to freshwater environments than Japan Sea sticklebacks. We suggest adaptive divergence between ancestral marine populations may have played a role in constraining phenotypic diversification and adaptive evolution in Japanese sticklebacks.     

Genetic evidence supporting the existence of two distinct species in the genusGasterosteus around Japan

Synopsis The genetic and morphological features ofGasterosteus aculeatus were investigated for 29 populations around Japan. Allozyme analyses recognized two groups (Pacific Ocean group and Japan Sea group) that had distinct characteristic features, and showed high genetic differentiation between them (D = 0.482). The Pacific Ocean group had a wide range, from North America to Japan, along the Pacific coast. The distribution of the Japan Sea group was limited around the Sea of Japan and the Sea of Okhotsk. The distribution of these groups were found to be sympatric on the Pacific coast of Hokkaido Island, Japan. From this area, genetic analyses demonstrated that the sympatric populations of the two groups formed independent breeding stocks, and it is considered that the two groups were reproductively isolated from each other. Additionally, each group had distinctive morphological features of lateral plates and caudal keels in the sympatric area. These results suggested that these two groups of the threespine stickleback comprise different species and that the Japan Sea group is taxonomically distinguishable fromG. aculeatus.     

Genetic structure of threespine stickleback,Gasterosteus aculeatus,in Lake Harutori,Japan, with reference to coexisting anadromous and freshwater forms

Both anadromous and freshwater forms of threespine sticklebacks (Gasterosteus aculeatus) inhabit Lake Harutori on the pacific coast of eastern Hokkaido, Japan. Since the two groups of threespine sticklebacks, Pacific Ocean and Japan Sea groups, which showed high genetic differentiation between them, were sympatrically distributed on the Pacific Ocean coast of Hokkaido Island, the genetic structures of the two forms were examined in Lake Harutori. Allozyme analyses of the two forms showed that most specimens from Lake Harutori belonged to the Pacific Ocean group, with a few fish belonging to the Japan Sea group or representing hybrids between the two groups. Both anadromous and freshwater sticklebacks were detected in the Pacific Ocean group. There were no significant differences in allelic frequencies at 17 presumptive loci between the two forms. Analyses of genetic relationships among 5 populations showed that the freshwater population from Lake Harutori was genetically more closely related to the sympatric anadromous population than to the freshwater population from a neighboring river. These results suggested that anadromous and freshwater sticklebacks in Lake Harutori form a single breeding population, and that the two forms might represent a life-history polymorphism within a single population.     

Extensive Introgression of Mitochondrial DNA Found between two Genetically Divergent Forms of Threespine Stickleback, Gasterosteus Aculeatus, around Japan

The genetic structure and phylogenetic relationships of Gasterosteus aculeatus around Japan were investigated for 15 populations of the Japan Sea and Pacific Ocean forms previously detected by allozyme analyses. PCR-RFLP analysis of mtDNA ND5/6 gene indicated two major divergent clades: (1) Alaskan population and (2) Russian and Japanese populations. Japanese populations were clearly subdivided into G. aculeatus populations and hariyo (G. aculeatus leiurus). However, the relationships among Japanese G. aculeatus populations did not support the genetic divergence between Japan Sea and Pacific Ocean forms shown by allozyme analyses. This suggests that mtDNA genetype similarity between the two forms could have resulted from complete replacement of mtDNA from one form to the other. MtDNA introgression is possible in two direction: from Japan Sea form to Pacific Ocean form and the reverse. The haplotype composition of each population observed in the present study suggests that the second direction is more likely. The migrant number of females per generation (Nm) between populations of the two forms was remarkablely large (9.083–infinite), suggesting that gene flow is taking place between the populations at the present time or might have occurred until recent years.     

SEGREGATIVE MIGRATION OF DALL'S PORPOISE (PHOCOENOIDES DALLI) IN THE SEA OF JAPAN AND SEA OF OKHOTSK

Dall's porpoises from the Sea of Japan population taken by a dolphin fishery vessel were examined. Estimated body lengths at sexual maturity were similar to those for the truei -type population off the Pacific coast of Japan but larger than those for the offshore dalli -type in the North Pacific. The Sea of Japan population is known to migrate to the Pacific coast of Japan through Tsugaru Strait and to the Sea of Okhotsk through Soya Strait in summer. From the difference of catch composition, individuals of different reproductive status in this population are considered to have different patterns of summer migration. Most of the individuals which migrate into the Pacific Ocean through Tsugaru Strait are sexually immature, while those migrating into the Sea of Okhotsk are mature. The Sea of Okhotsk is thought to be a breeding area for this population. The calving season is estimated to be May through June, which is earlier than that of the truei -type off the Pacific coast of northern Japan. This may be an adaptation to lower water temperatures in the Sea of Japan during winter.     

Bioaccumulation of lithium by marine organisms in European, American, and Asian coastal zones: microanalytic study using secondary ion emission

37 species of aquatic mammals, fish, crustacea, annelids, molluscs belonging to cephalopods, gasteropods and lamellibranchs were collected from coastal waters of France: North Sea, English Channel, Atlantic Ocean, Mediterranean Sea, from Greece: Aegian Sea, from North America: Atlantic Ocean and from Japan: Pacific Ocean, Sea of Japan and East China Sea. Microanalyses which were performed on organs and tissues, using secondary ion mass spectrometry, revealed high concentrations of lithium, which is commonly used in human therapy, but is also toxic in low amounts. The retention of this metal by the marine organisms appears as a general phenomenon independent of their biotope and geographical origin; the highest lithium levels were detected in the fish muscles (= edible part).     

Offshore Dolly Varden charr (<Emphasis Type="Italic">Salvelinus malma</Emphasis>) in the North Pacific

Many studies have investigated the ecology of charrs in freshwater, however, little is known about charrs in the ocean. This study examined the distribution, seasonal abundance, and some biological features of Dolly Varden (Salvelinus malma) in the Pacific Ocean. An analysis of by-catch data of Japanese offshore salmon monitoring showed that Dolly Varden were distributed across a wide range in the offshore waters of the Pacific Ocean, including the Japan Sea, Bering Sea, and Okhotsk Sea. The catch per unit effort showed a sharp increase from May to August, followed by a sharp decrease in September. Offshore areas served as an important summer habitat for anadromous Dolly Varden.     

Holoplankton of ship ballast water in the Port of Vladivostok

Ballast water samples were taken from three ships that arrived at the Port of Vladivostok (Peter the Great Bay, Sea of Japan) from ports of Japan (Sea of Japan and Pacific Ocean) and China (Yellow Sea and Yangtze River). The holoplankton in samples was presented by seven taxonomic groups, among which copepods (subclass Copepoda, 33 species) and cladocerans (subclass Cladocera, 5 species) dominated. In the samples, eight nonindigenous copepod species were revealed. The information contained in this paper may be of importance for the assessment of the risk of subsequent invasion of new species and the development of techniques for monitoring the qualitative and quantitative structures of ballast water.     

Distinct spawning migration patterns in sympatric Japan Sea and Pacific Ocean forms of threespine stickleback Gasterosteus aculeatus

Spawning migration patterns were compared between sympatric populations of the anadromous Japan Sea (JS) form and Pacific Ocean (PO) form of the threespine stickleback Gasterosteus aculeatus in the Lake Akkeshi–Bekanbeushi River system, eastern Hokkaido, Japan. In Akkeshi Bay and Lake Akkeshi, the JS form had longer collecting peak periods and more collection peaks than the PO form. Furthermore, although the PO form migrated upstream in freshwater habitats, the greater part of the JS form stayed in brackish waters before breeding. These findings suggest that the 2 forms have distinctly different spawning migration patterns and breeding grounds.     

GEOGRAPHIC VARIATION AND SEXUAL DIMORPHISM IN THE SKULL OF DALL'S PORPOISE, PHOCOENOIDES DALLI

Abstract: Two hundred and eighty-nine skulls of Dall's porpoises from the North Pacific Ocean, Sea of Japan, Sea of Okhotsk, and Bering Sea were investigated morphometrically. A clear pattern of geographic variation was found in overall size of the skull. In the North Pacific, overall skull size became gradually smaller eastward from the coast of Japan to the offshore eastern Pacific, and it became larger again off the coast of California. Specimens from the Sea of Japan-Okhotsk and Bering Sea had larger skulls than those from the central North Pacific. The distribution of primary productivity corresponds well with this pattern of geographic variation, suggesting that quantity of food might affect the overall size of Dall's porpoise skulls. By canonical discriminant analyses, about 70% (male) and 90% (female) of specimens of the Sea of Japan-Okhotsk population were distinguished from those of other populations. This agrees with the results of genetic studies.     

A new species of Enterognathus (Copepoda, Cyclopoida, Enterognathidae) collected from the Seto Inland Sea, western Japan

A new species of the endoparasitic copepod Enterognathus (Cyclopoida, Enterognathidae) is described from a crinoid host in the Seto Inland Sea, western Japan. This is a third species of the genus and its first occurrence in the Pacific Ocean. The new species is distinguished from two previously known congeners by the morphology of the body somites, caudal rami, antennae and legs. Crinoid parasites belonging to Enterognathus and the closely related genus Parenterognathus have a broad distribution from the northeastern Atlantic through the Red Sea to the West Pacific.     

A concise review of geographic variation in adult body size in anadromous masu salmon, Oncorhynchus masou

Adult body size (size at maturity) is one of the key life history traits and is well known to sometimes correlate with latitude in anadromous salmonids. However, it is poorly understood whether geographic size patterns except for latitudinal trends occur or why such patterns have been shaped. The present paper briefly reviewed body size variation between anadromous returns of masu salmon Oncorhynchus masou in the Okhotsk group (10 populations along the Sea of Okhotsk coast), the Pacific group (2 populations along the Pacific Ocean coast) and the Sea of Japan group (24 populations along the Sea of Japan coast). The Okhotsk group was smaller than the Sea of Japan group. Although the statistical analysis detected no differences among the remaining combinations, the Okhotsk group was possibly smaller than the other groups because the size of the Pacific group seemed to be within range of the Sea of Japan group but tended to be larger than that of the Okhotsk group. Future research should first test whether size at maturity genetically differs between the Okhotsk group and the other two groups to explore further evolutionary factors shaping geographic size variation.     

Genetic structure of the beaked whale genus Berardius in the North Pacific,with genetic evidence for a new species

There are two recognized species in the genus Berardius, Baird's and Arnoux's beaked whales. In Japan, whalers have traditionally recognized two forms of Baird's beaked whales, the common “slate‐gray” form and a smaller, rare “black” form. Previous comparison of mtDNA control region sequences from three black specimens to gray specimens around Japan indicated that the two forms comprise different stocks and potentially different species. We have expanded sampling to include control region haplotypes of 178 Baird's beaked whales from across their range in the North Pacific. We identified five additional specimens of the black form from the Aleutian Islands and Bering Sea, for a total of eight “black” specimens. The divergence between mtDNA haplotypes of the black and gray forms of Baird's beaked whale was greater than their divergence from the congeneric Arnoux's beaked whale found in the Southern Ocean, and similar to that observed among other congeneric beaked whale species. Taken together, genetic evidence from specimens in Japan and across the North Pacific, combined with evidence of smaller adult body size, indicate presence of an unnamed species of Berardius in the North Pacific.     

Population Structure of Pacific Cod <Emphasis Type="Italic">Gadus macrocephalus</Emphasis> in the Southern Part of the Range Based on the Microsatellite Analyses

The population structure of Pacific cod Gadus macrocephalus in the southern part of the range and adjacent regions is studied on the basis of the results of microsatellite analyses. Collected data indicate heterogeneity of this species population within the studied area. According to the obtained F_ST values, Pacific cod from the waters of the Republic of Korea (Yellow Sea side) and northwestern part of the Sea of Okhotsk significantly differ from all other studied regions (Table 4). Significant differentiation was also revealed between samples from the waters of the Tatar Strait and all other regions except for South Kurils Pacific cod (both Sea of Okhotsk and Pacific Ocean sides). These two latter sample collections were similar to each other as well. A low level of differentiation was also shown for the Peter the Great Bay and the East Sea/Sea of Japan waters of the Republic of Korea.     

Detection of regional allozyme divergence in the rocky intertidal goby Chaenogobius annularis

Genetic differentiation in the intertidal goby Chaenogobius annularis was studied using allozyme markers. Samples were collected from six localities along the coasts of Japanese islands and the Korean Peninsula. Six out of 13 loci showed allelic variation in at least one population. Although the predominant alleles of 12 loci were the same among all populations, Mdh-1 showed clear differences among the populations located along the coasts of the Pacific Ocean and the Sea of Japan. These two possible geographic groups, the Pacific Ocean and the Sea of Japan groups, were characterized by diagnostic alleles of Mdh-1, namely Mdh-1 ¹⁰⁰ and Mdh-1 ⁷⁰.     

Phylogeography and historical demography of the anadromous fish Leucopsarion petersii in relation to geological history and oceanography around the Japanese Archipelago

Phylogeographical patterns of marine and diadromous organisms are often influenced by dynamic ocean histories. For example, the marine realm around the Japanese Archipelago is an interesting area for phylogeographical research because of the wide variation in the environments driven by repeated shifts in sea level in the Quaternary. We analysed mitochondrial cyt b gene and nuclear myh6 gene sequences for individuals collected from throughout the range of the anadromous fish Leucopsarion petersii to assess the lineage divergence, phylogeographical pattern and historical demography in relation to geological history and oceanographic features around the archipelago. Leucopsarion petersii has two major lineages (the Japan Sea and Pacific Ocean lineages), which diverged during the late-early to middle Pleistocene. Geographical distributions of the two lineages were closely related to the pathways of the two warm currents, the Tsushima Current and the Kuroshio Current, that flow past the archipelago. Evidence of introgressive hybridization between these lineages was found at two secondary contact zones. Demographic tests suggested that the Japan Sea and Pacific Ocean lineages carried the genetic signal of different historical demographic processes, and these signals are probably associated with differences in habitat stability during recent glacial periods. The Japan Sea lineage has a larger body-size and more vertebrae, probably in relation to severe habitat conditions through Pleistocene climatic oscillations. Thus, the two lineages have long independent evolutionary histories, and the phylogeographical structure and demography of this species have been influenced both by historical events and the present-day oceanography around the Japanese Archipelago.     

Mackerel from the northern indian ocean and the red sea arescomber australasicus, notscomber japonicus

The population ofScomber from the Red Sea and northern Indian Ocean (gulfs of Aden and Oman) is identified asS. australasicus rather thanS. japonicus based on having 30–33 vs. 26–29 interneural bones under the first and second dorsal fins and the combination of interneural bone counts of 16–20 under the first dorsal fin (vs. 13–16) and first dorsal fin spine counts of 10–13 (vs. 9–10). These are the best morphological characters to distinguish these two species. This change in identification constitutes a major range extension forS. australasicus which was thought to be restricted to the Pacific Ocean and the southeastern Indian Ocean around Western Australia.     

Evidence for initial opening of the Pacific ocean in the Jurassic

An argument is presented for the east-west initial opening of the pacific Ocean during Early Jurassic times, on paleontological, biogeograhical, and geological grounds. A new reconstruction of the continents bordering the Pacific Ocean is proposed, based chiefly upon trans-Pacific, terrestrial biotic links that are not reflected in regions bordering the Atlantic. A new Arctic Ocean reconstruction is also proposed. Since the Atlantic and Indian Oceans were closed in the Early Jurassic as well, an expanding-earth model is adopted, instead of a Pangaen reconstruction on a present-sized earth. The model prposed here is also compatible with geological evidence for a narrow, epicontinental Tethys Sea. Sea-floor subduction along the perimeter of the Pacific appears to be actively taking place and in the past was much more extensive along the eastern border than on the western side. The tectonics of Antarctica, the Southwest Pacific, Indonesia, Japan, the Arctic region, western North America, Baja California, the Carribean, and the Gulf of Mexico are discussed in terms of the proposed model. The Pacific sea-floor spreading history is outlined and related to the initial opening of the North Atlantic 180 million years ago (Early Jurassic). The South Pacific and Indian Ocean are both considered to have opened some 155 million years ago (Late Jurassic). The original and present positions of the Jurassic equator are determined.