Comments on cephalometrical configurations of the Japanese

Cephalometrical configurations of Japanese people were investigated on two hundred and sixteen groups living on the four main islands of Japan: Hokkaido, Honshu, Shikoku, and Kyushu. The Japanese in Honshu were divided geographically into a narrow-featured eastern major group and a broad-featured western major group. A physical characteristic of Japanese living in the western mid-Honshu was the lengthened feature of their cephalic measurement, an influence of a long-headed group of Korean people; Japanese residents in eastern mid-Honshu, however, showed the result of mixing with a short-headed group of Korean people. Japanese people living in Hokkaido, the northernmost island, and in the Ryukyu Islands in the South of Japan, presented no important finding which suggests their influence on the physical characteristics of the Japanese. Ongoing studies indicating physical relationships between Kyushu or Shikoku residents and Honshu residents, are as yet insufficient.     

A new timeframe for the diversification of Japan’s mammals

Aim The main Japanese islands are land‐bridge islands divided by the biogeographic division Blakiston’s Line and represent two natural laboratories for studying land‐bridge diversification. Colonization of the current mammal fauna has been dated to the middle to late Pleistocene using fossil evidence. The purpose of this paper is to apply a molecular clock to the genetic divergences between Japanese mammalian taxa and their sister mainland taxa to test the late Pleistocene land‐bridge colonization hypothesis. Location The main Japanese islands (Kyushu, Shikoku, Honshu and Hokkaido). Methods I used mitochondrial DNA (cytochrome b) and a species tree approach to estimate the divergence times of 24 Japanese non‐volant terrestrial mammal taxa and their mainland sister taxa using the program *beast . I then tested for evidence of non‐simultaneous divergence among these taxon‐pairs by controlling for expected coalescent stochasticity using the program Ms Bayes . Results Divergence events between taxa on Japan and their mainland sister taxa were significantly older than expected under the current paradigm, which is based on fossil data. Consistent with the land‐bridge colonization hypothesis, there was evidence of multiple divergence events. Main conclusions These results implicate a colonization timeframe that is older than posited by the current paradigm based on fossil evidence. However, these results are still consistent with the land‐bridge colonization hypothesis. Multiple periods of land‐bridge connectivity may account for the current mammalian fauna in Japan. In addition, half of the divergence time estimates in the Honshu–Shikoku–Kyushu region were clumped around 2.4 Ma, which might suggest a dramatic interchange period, concordant with a period of significant global cooling, when the first land bridge may have connected Japan to the mainland.     

Differential geographic patterns of mitochondrial DNA variation in two sympatric species of Japanese wood mice, Apodemus speciosus and A. argenteus

We examined the gene sequences of mitochondrial cytochrome b (cyt b) in two Japanese wood mouse species, Apodemus speciosus (n = 89) and A. argenteus (n = 46), which are distributed on the four main islands of Japan (Hokkaido, Honshu, Shikoku, and Kyushu) and on the small islands surrounding them. Apodemus speciosus, the larger of the two species, showed substantial genetic variation, with a maximum of 3% sequence divergence, and remarkable phylogenetic subdivision with two major clades. One clade represents haplotypes from a central region, including Honshu, Shikoku, Kyushu, and their adjacent islands; the other clade includes haplotypes from Hokkaido and the peripheral islands, forming four subclades: a) Hokkaido, b) Sado Island, c) Satsunan Islands, and d) the Izu Islands. Sequence divergence among the four subclades was 1.0 to 1.5%, implying that A. speciosus colonized these geographic regions 0.2 to 0.3 million years ago, assuming a substitution rate of 2.4% per million years. The population on the Izu Islands has preserved haplotypes that are distinct from those in any other region, providing good evidence for the natural colonization of the volcanic islands of the Izu Islands. The cyt b sequence variation had no relation to the karyotypic dimorphism for the eastern (2n = 48) and western (2n = 46) geographic groups, between which a strict border exists at central Honshu. On the other hand, Apodemus argenteus, the smaller of the two species, showed a similar level of sequence divergence (maximum of 3%) but no substantial geographic differentiation: populations in Hokkaido, Sado, and Yakushima shared similar haplotypes with each of the central populations, suggesting that genetic exchanges occurred among the localities in the last 0.15 million years. The apparent genetic structure of the mitochondrial DNA found in the A. speciosus population might be caused solely by long-term existence in insular regions, presumably due to ecological superiority relative to A. argenteus.     

Prehistoric Sado Island populations of Sus scrofa distinguished from contemporary Japanese wild boar by ancient mitochondrial DNA

Ancient mitochondrial DNA (mtDNA) mainly from Jomon Period Sus scrofa bone specimens (6,100-1,700 years old) was examined to clarify the genetic relationships between prehistoric and contemporary S. scrofa on Hokkaido, Honshu, Sado, and Izu islands of the Japanese Archipelago. Phylogenetic analysis of the mtDNA control region (574 bp) and analysis of pairwise nucleotide differences between prehistoric and contemporary S. scrofa sequences showed the following relationships between these groups: (1) a group genetically similar to contemporary Japanese wild boars was found mainly on Honshu Island, Hokkaido Island, and the Izu Islands, and (2) a monophyletic group distinct from contemporary Japanese wild boars was found on Sado Island. These results suggest that prehistoric people introduced S. scrofa from Honshu Island to Hokkaido Island and the Izu Islands. The estimated divergence times between the prehistoric Sado group and the other prehistoric S. scrofa is approximately congruent with the geological isolation of Sado Island from Honshu Island. Our results suggest that this extinct S. scrofa population was present on Sado Island as recently as around 2,000 years ago.     

Phylogeography of Japanese population of Phelotrupes auratus (Coleoptera, Geotrupidae) inferred from mitochondrial DNA sequences

A partial sequence of the mitochondrial cytochrome c oxidase subunit I (COI) gene (745 bp) was determined for 57 specimens of a geotrupid beetle (Phelotrupes auratus) from throughout the Japanese archipelago. Of the 57 beetles examined, 42 haplotypes were identified. Phylogenetic trees inferred using maximum parsimony, neighbor joining, and Bayesian inference methods were highly congruent. Reconstructed phylogenetic relationships indicated that P. auratus from the Japanese archipelago was separated into two distinct lineages: Group A, which consisted of 35 haplotypes from Honshu, Shikoku, and Hokkaido Islands, and Group B, which consisted of seven haplotypes from Kyushu and Yakushima Islands. In addition, two sublineages were also recognized within Group A: Subgroup A-1, which consisted of 11 haplotypes from eastern Honshu and Hokkaido, and Subgroup A-2, which consisted of 10 haplotypes from western Honshu and Shikoku. Average genetic distances within Group A were positively correlated to geographic distance between sampling localities. Phylogenetic relationships among haplotypes did not correspond to subspecies classification.     

Mitochondrial DNA phylogeography of the red fox (Vulpes vulpes) in northern Japan

Mitochondrial DNA variation in the cytochrome b (cyt b) gene and the control region was examined in the red fox Vulpes vulpes from Japan, with special focus on the population divergence between Hokkaido and northern Honshu. Resultant haplotypes from Hokkaido were subdivided into two distinct groups (I and II), with an average genetic distance of 0.027 for cyt b. Divergence time is roughly estimated to be 1-2 million years ago, given that the conventional divergence rate of the mammalian cyt b gene is 2% per million years. Notably, Group II was only found in Hokkaido, whereas Group I comprised haplotypes from Honshu, Kyushu (Japan), eastern Russia, and Europe, as indicated by a comparison of our own data to the literature. On the other hand, judging from constructed trees, Group I haplotypes from Hokkaido appeared to differ from those from other parts of Japan, i.e., Honshu and Kyushu. This implies that Blakiston's Line, which demarcates the boundary between Hokkaido and Honshu, has been an effective barrier and has allowed the structuring of genetic variation in maternal lineages. Thus, these results suggest that the Hokkaido population, which is sometimes referred to as the distinct subspecies V. v. schrencki, has its own genetic background with multiple migration events and differs from the parapatric subspecies V. v. japonica found in Honshu and Kyushu.     

Molecular phylogeny of the genus Apatrobus (Coleoptera: Carabidae: Patrobinae) in western Japan

We performed a molecular phylogenetic analysis of the ground beetles Apatrobus (Carabidae), endemic to Japan, using the mitochondrial cytochrome c oxidase subunit I (COI) and the nuclear 28S rRNA (28S) genes. We focused on the species divergence in Kyushu, Shikoku and western Honshu and used 15 of 19 species and three populations with undetermined species in the DNA analysis. The gene trees showed that, of the Apatrobus species studied, A. hayachinensis Nakane from northern Honshu was not included in the monophyletic group of the other Apatrobus species and likely to be of a different genus. Divergence time estimation suggested that Apatrobus species excluding A. hayachinensis diverged 5.2 million years ago and the subsequent divergence of species occurred during the Pliocene and Pleistocene. In each of the main islands, Kyushu, Shikoku and Honshu, two or more distinct lineages occurred and all species had restricted distribution areas, suggesting that ancient dispersal and vicariance among the three main islands resulted in the nested biogeographical pattern of species distribution.     

The genus Campylocheta (Diptera: Tachinidae) from Japan

Seven new species of Campylocheta are described and illustrated from Japan: abdominalis from Hokkaido, Honshu and Kyushu, flaviceps from Honshu and Tsushima, grisea from Kyushu, argenticeps from Honshu, bisetosa from Hokkaido and Honshu, hirticeps from Hokkaido, Honshu and Tsushima, and suwai from Hokkaido. Campylocheta dentifera Richter and C. umbrinervis Mesnil are redescribed and a key to nine Japanese species is provided.     

Spatial framework of nine distinct local populations of the Japanese dormouse Glirulus japonicus based on matrilineal cytochrome b and patrilineal SRY gene sequences

We previously revealed the presence of six genetically distinct matrilineal populations of the Japanese dormouse Glirulus japonicus in the distribution range of Honshu, Shikoku, and Kyushu islands. In this study, we extended this analysis using mitochondrial cytochrome b gene sequences (n = 96) and Y-chromosome-specific SRY gene sequences (n = 22) from individuals collected from Honshu, Shikoku, Kyushu, and Oki Dogo I. The cytochrome b sequence data allowed us to define precise geographic ranges of the six previously known and three newly found distinct matrilineal lineages: northeastern Honshu (I), east-central Honshu (II), west-central Honshu and the Kii Peninsula (III), the western part of Honshu (IV), Shikoku (V), westernmost Honshu and Kyushu (VI), the northern part of central Honshu (VII), the southern part of central Honshu (VIII), and Oki Dogo I. (IX). Our inference of geographic borders suggests that regions of lower and higher altitudes in the mountain systems played important roles in driving the hosting and separation of lineages, respectively. Six matrilineal lineages (I, II, V, VI, VIII, and XI) were shown to possess their own SRY haplotypes, while lineages III and IV shared one haplotype. These data together with our previous observation of nuclear ribosomal RNA gene variation indicate advanced populational subdivision in this species. It is thus evident that each of the populations, including those living at high latitudes and in limited geographic spaces, have survived for several million years. A specific ability to tolerate cold may have permitted G. japonicus to preserve anciently diverged lineages in each locality.     

Two mitochondrial lineages occur in the Asian corn borer, Ostrinia furnacalis (Lepidoptera: Crambidae), in Japan

The genealogy and diversity of the mitochondrial cytochrome oxidase subunit II (COII) gene were investigated for Ostrinia furnacalis in Japan. A preliminary examination of mitochondrial lineages in China and the Philippines was also made. Two lineages (A and B) were found in the COII gene. Lineage A was frequent throughout the Japanese main islands (Hokkaido, Honshu, Shikoku and Kyushu), while the frequency of lineage B varied among these islands. No clear patterns of geographical population structure were found. Population genetic features suggested that the O. furnacalis population harboring the lineage A mitochondria expanded in the recent past, while lineage B showed weak signals of a population expansion. It is not clear whether the two lineages of mtDNA evolved in separate or identical geographical regions. We discuss two hypotheses regarding the two lineages of mtDNA: a cryptic race/species hypothesis and a selective sweep hypothesis.     

Mitochondrial DNA diversification, molecular phylogeny, and biogeography of the primitive rhacophorid genus Buergeria in East Asia

In this study we sought to clarify the evolutionary relationships and biogeographic history of the bell-ring frog, Buergeria buergeri (family Rhacophoridae), and two congeneric species Buergeria japonica and Buergeria robusta, by analyzing three mitochondrial (mt) genes: 12S rRNA, Cytb, and ND5. Phylogenetic analyses based on gene data showed the mt clades corresponding to the Buergeria species and three major haplogroups within B. buergeri. Each haplogroup corresponded clearly to the area in which it was distributed, namely eastern Japan (excluding Hokkaido; Hg I), central Japan (Hg II), and western Japan (including the Shikoku and Kyushu regions; Hg III). The estimated divergence time suggested that the lineage splits of the Buergeria species occurred before the formation of the island of Taiwan and the Japan Archipelago (including the Ryukyu islands). The differentiation among the genealogical lineages of B. buergeri seems to have begun in the Late Miocene (approx. 7-5Mya), and the formation of their present distribution pattern might have been influenced by climatic changes and geographical events such as the formation of a wide peneplane and expansions of certain basins.     

Phylogeography and population structure of the Japanese wild boar Sus scrofa leucomystax: mitochondrial DNA variation

Phylogeographic characteristics and population structure of Japanese wild boar (Sus scrofa leucomystax) were investigated using mitochondrial DNA (mtDNA) sequence data. Sixteen Japanese wild boar haplotypes detected from partial sequences of the mtDNA control region (574-bp) from 180 Japanese wild boar specimens from 10 local populations on Honshu, Shikoku, and Kyushu islands and 41 haplotypes from other S. scrofa were analyzed using the neighbor-joining method. The Japanese wild boars were more closely related to Northeast Asian wild boars from Mongolia than to the other Asian continental S. scrofa. The Japanese and Northeast Asian wild boars were not significantly distinguished by corrected average pairwise difference analysis. The ancestors of Japanese wild boars are suggested to have been part of the continental S. scrofa population that spread from Southeast to Northeast Asia during the Middle to Late Pleistocene. The Japanese wild boar mtDNA haplotype cladogram shows 95% parsimoniously plausible branch connections supporting three sympatric clades. Nested clade analysis indicates that these three clades are the result of distinct historical events or gene flow. The present population of Japanese wild boars may have been formed by a few independent migrations of distinct clades from the continent with subsequent mixing on the Japanese Islands.     

Community composition and size structure of murid rodents in relation to the biogeography of the Japanese archipelago

We investigated the geographical patterns of community composition and size structure of murid rodent assemblages in Japan. Rodent faunal composition showed three biogeograpbic zones in the studied area (Hokkaido, northern Honshu and southern Honshu), which are characterized by endemic species or genera. There was a large discrepancy between distribution patterns of murine species, which are generalist and widespread in Japan, and arvicoline species, which are more specialized and locally restricted. We also found a strong degree of nestedness of the murid rodent fauna, i.e. smaller faunas were subsets of larger ones, which is typical of relict fauna. The structure of murid rodent assemblages was studied using the size and shape of the lower incisor, in order to test for the effect of interspecific competition on community-wide patterns. We used two different approaches: one tests for regularity in the size structure of the community (Barton and David test), and the other one tests for minimum mean overlap size in the community between species (randomization procedure). There was no congruence between the results of the two tests: we did not find any case of regular size structure, whereas mean size overlaps were minimum or even zero in about half of the cases studied. Thus, the evolution of rodent communities on islands seems to be characterized by minimization of size overlaps, perhaps as a result of interspecific competition. Also, the reduction of island area, which is correlated with a decrease in species richness, is accompanied by an increase in Hutchinsonian size ratios and a decrease in the total size range of the community. These patterns may be linked to the reduced diversity of environmental resources on islands.     

Geographic allozyme variation in the Japanese clouded salamander, Hynobius nebulosus (Amphibia: Urodela)

Using starch gel electrophoresis, geographical enzyme variation was surveyed in 46 populations of the Japanese salamander Hynobius nebulosus , which occurs widely in western Japan. This species exhibits substantial local genetic differentiation and is more diverse in inland regions than on small islands. In several different analyses, two groups of populations, one from Kyushu Island to the westernmost part of Chugoku district, Honshu (western group), and another from Shikoku through Kinki to Chubu district, Honshu (eastern group), were consistently recognized. Most of the remaining populations were placed in two less clearly-defined groups: the montane group from the Chugoku Mountains and the Chugoku group from the coastal regions of Chugoku district. The divergence patterns of H. nebulosus are thought to be related to the geological history of its range. The populations near the boundary of the western group were found to share several alleles that were predominant on both sides of the boundary, indicating past secondary contact. Differentiation in the montane group did not follow the isolation by distance model, probably because several populations included were genetically mixed with lowland populations of the Chugoku group.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 311–330.     

Onset of cryptic vicariance in the Japanese dormouse Glirulus japonicus (Mammalia, Rodentia) in the Late Tertiary, inferred from mitochondrial and nuclear DNA analysis

The sequences of the mitochondrial cytochrome b gene and restriction site variation in the spacer region of the nuclear ribosomal RNA gene [rDNA-restriction fragment length polymorphism (RFLP)] were analysed to determine the phylogeographic structure of the Japanese dormouse ( Glirulus japonicus ), which is threatened by deforestation and has been designated an endangered species in Japan. The phylogenetic tree of cytochrome b grouped G. japonicus into six geographical populations: north-eastern Honshu (I), central Honshu (II), west-central Honshu/Kii Peninsula (III), western Honshu (IV), Shikoku (V), and westernmost Honshu/Kyushu (VI); the genetic distances among these groups suggest divergence in the Late Tertiary. The lineage of group VI was located at the basal position in the phylogenetic tree, followed by the radiation of the other lineages. An rDNA-RFLP analysis of 15 restriction sites roughly supported such genetic isolation; groups I, II, III, IV, V and VI have five, two, one, one, one and four unique restriction sites, respectively, revealing four geographic groups as cryptic species: I, II, III + IV + V and VI. Our results reveal the ancient divergences of the local population, which has a complicated evolutionary history, and should be useful in developing a framework for the conservation of this species.     

Patterns of distribution in Japanese land mammals

Distribution patterns among the terrestrial mammal species of Sakhalin and the main islands of Japan are shown to fall into 12 clear groups. The most fundamental distributional break (Blakiston's Line) is that separating Hokkaido and Sakhalin to the north, with their boreal fauna typical of northern Eurasia, from ‘Hondo’ (Honshu, Shikoku, Kyushu) to the south, which demonstrates a high degree of endemism and supports a small number of Indo-Malayan elements. Distribution patterns may be explained by considering the Quaternary geohistory of the area, particularly the formation of land bridges and the changes in climatic conditions during this period. A dynamic classification based upon origin of the fauna recognizes seven categories, although on evidence currently available the allocation of some species to categories is not yet possible. Hondo underwent two main periods of land bridge connection to the Asiatic mainland. The first, prior to the Pleistocene, allowed immigration of forms which have since developed into distinctive elements of the endemic fauna (‘Old Hondo Endemics’). The second, during the Middle Pleistocene, brought in widespread Palaearctic species as well as components from South-East Asia (‘Early Colonists’), some of which have since undergone vicariant speciation (‘New Hondo Endemics’); it also allowed several of the Old Hondo Endemics to extend their range to the mainland and Hokkaido (‘Expanding Hondo Endemics’). Sakhalin and Hokkaido have been more intimately connected to the mainland (most recently until less than 10 000 years ago), such that endemism is very restricted. Species groups here are the ‘Late Colonists’, cold-adapted tundra species which expanded with the glacial advances, but which are now restricted in distribution, and ‘Recent Colonists’, postglacial forest species which recolonized before the severance of land bridges. Moving the other way were ‘Expanding Northern Endemics’, which arose in Hokkaido or Sakhalin during the last glacial and colonized the adjacent mainland before severance of land links. There are, additionally, several possible Expanding Hondo Endemics in Hokkaido, although human intervention in determining their current distributions cannot be ruled out. Bats, to which the sea barriers of the Japanese area are less complete, exhibit somewhat different patterns of distribution, confirming predictions about the role of even narrow marine straits in restricting ranges of nonflying mammals.     

Molecular phylogeny of Japanese Leporidae,the Amami rabbit Pentalagus furnessi,the Japanese hare Lepus brachyurus,and the mountain hare Lepus timidus,inferred from mitochondrial DNA sequences

We determined mitochondrial 12S ribosomal RNA (rRNA) and cytochrome b (cyt b) gene sequences in three leporid species of Japan, the Amami rabbit Pentalagus furnessi from the Ryukyu Islands, the Japanese hare Lepus brachyurus from Honshu, and a Japanese form of the mountain hare Lepus timidus ainu from Hokkaido. We compared the sequences with those of other taxa of leporids available in databases. Phylogenetic trees of the 12S rRNA gene sequences indicated that the lineage of P. furnessi diversified during the generic radiation of the leporids at an ancient time, which was estimated to have been the middle Miocene. Cyt-b gene trees revealed that the lineage of L. brachyurus branched off at an early stage in the speciation of Lepus, probably at the beginning of the Pliocene. The cyt b sequences of L. t. ainu were somewhat distinct from those of continental conspecific populations; this lineage divergence is likely to have occurred during the middle or late Pleistocene. The results show that the three regions of the Japanese archipelago, Ryukyu, Honshu-Shikoku-Kyushu, and Hokkaido, now preserve their own leporid taxa, each with a different extent of genetic endemicity. It is possible that the zoogeographic traits of the Japanese leporids are a consequence of the evolutionary dynamics of leporids in East Asia, in that the radiation centers of leporids are likely to have shifted from tropical, through temperate, to arctic zones.     

Distribution of the four flagellar (H) antigenic subserotypes of Bacillus thuringiensis H serotype 3 in Japan

A total of 302 strains of Bacillus thuringiensis flagellar (H) serotype 3, collected from sericultural environments and soils of the three main islands (Honshu, Shikoku and Kyushu) of Japan, were examined for their H antigenic subserotypes. Of these strains, 259 (85.8%) were identified as the H subserotype 3a : 3c (subsp. alesti ), 16 (5.3%) were referable to the subserotype 3a : 3b : 3c (subsp. kurstaki ), 26 (8.6%) belonged to the subserotype 3a : 3d : 3e (subsp. fukuokaensis ) and one strain (0.3%) was the subserotype 3a : 3d (subsp. sumiyoshiensis ). The subserotypes 3a : 3c and 3a : 3b : 3c were present in sericultural environments only; the former was distributed in Honshu and Shikoku Islands but not in Kyushu Island, and the latter was distributed in Honshu and Kyushu Islands but not in Shikoku Island. The subserotype 3a : 3d : 3e, mainly found in soils but occasionally in sericultural environments, was distributed in the three main islands. It also appeared that the subserotype 3a : 3d : 3e, a recently described subserotype, was distributed in Japan more widely than the globally disseminated subserotype 3a : 3b : 3c. The results clearly showed geographical and ecological differences in the distribution of subserotypes.     

A trend of central versus peripheral structuring in mitochondrial and nuclear gene sequences of the Japanese wood mouse, Apodemus speciosus

A phylogeographic analysis was performed on Japanese endemic wood mice (Apodemus speciosus) using nuclear interphotoreceptor retinol binding protein (IRBP) gene sequences (1,152 bp), together with previously published mitochondrial cytochrome b (cyt b) data. In the IRBP analysis, 40 haplotypes were recovered from 84 individuals by statistical and subcloning methods. Substantial sequence variation was determined from the IRBP data (pi=0.0047), and no significant evidence of recombination was detected. From the phylogenetic analysis, the 40 haplotypes fell into two major groups with geographic associations, irrespective of the karyotype groups (2n=46 and 2n=48), yielding a trend of central (Hokkaido, Honshu, Shikoku, Kyushu, and Sado) and peripheral (Izu, Oki, Tsushima, and Satsunan Is.) groupings. This geographic pattern is similar to that observed in the cyt b data, with a different insular grouping of Sado, Hokkaido, Izu, and Satsunan islands, and also to that of morphological features. In both gene data sets, nested clade analyses revealed allopatric fragmentation in the "peripheral island clades" and range expansion in the "central island clades." A mismatch analysis using cyt b data also suggested expansion of the central islands clade. Thus, the trend of central vs. peripheral structuring may be attributable to past demographic dynamics in the two distinct haplotype clades, such as range expansion of one clade in the central area of the Japanese Islands, leaving the other clade in the periphery.