Climatic controls on distribution of Fagus crenata forests in Japan

Abstract. We used classification tree analysis to develop a climate‐based distribution model for Fagus crenata forests in Japan. Four climatic variables judged likely to affect the distribution of the species (summer and winter precipitation, minimum temperature of the coldest month and Kira's warmth index) were chosen as independent variables for the model. Latitudinal and longitudinal information was also used to examine effects of spatial autocorrelation on the model. The climatic factors associated with the distribution of the forests were analysed using a classification tree to devise prediction rules. Predicted areas of high probability for forest occurrence lay mainly on the Sea of Japan side of northern Honshu and southern Hokkaido. This is consistent with actual forest distribution. Some areas with high predicted probabilities of F. crenata forest occurrence were beyond the current natural northern range limits of these forests. Since these areas were widely scattered, it was assumed that the species has been hindered from colonizing them due to dispersal limitations. Deviance‐weighted scores, used to compare magnitudes of the contributions of predictor variables, revealed winter precipitation as the most influential factor, followed by the warmth index, the minimum temperature of the coldest month and summer precipitation. Attempts were made to generate ecological explanations for the effects of the four climatic factors on the distribution of F. crenata forests.     

Ploidy, geographical distribution and morphological differentiation of Parasenecio auriculata (Senecioneae; Asteraceae) in Japan

Parasenecio auriculata is a woodland perennial herb widely distributed in Northeastern Asia, constituted by a poorly understood polyploidy with a diploid (2n=2x=60) and a tetraploid (2n=4x=120). In this study, for a better understanding of the polyploidal evolution, cytogeography and morphological variation were analyzed in Japanese P. auriculata, including two varieties; var. bulbifera endemic to central Hokkaido and var. kamtschatica widely distributed in northern Honshu and Hokkaido. The occurrence of two polyploidal levels was reconfirmed. While var. bulbifera is predominantly tetraploid, var. kamtschatica is comprised of diploid and tetraploid. Morphological variation among 22 quantitative characteristics is continuous and not distinctive among cytotypes or varieties, but plant size tended to be larger in the order, diploid of var. kamtschatica, var. bulbifera, and tetraploid of var. kamtschatica. The cytotype distribution showed a conspicuous geographical pattern. Besides var. bulbifera endemic to the central Hokkaido, the diploid of var. kamtschatica is mainly found in Southern Hokkaido, and the tetraploid has a disjunct distribution in eastern and northern Hokkaido and northern Honshu. Such a geographical pattern is possibly attributable to the differentiation of climatic preference among cytotypes and varieties, and may have been established in association with the climatic cline along the Japanese archipelago.     

Different phylogeographic patterns in two Japanese Silpha species (Coleoptera: Silphidae) affected by climatic gradients and topography

To reveal differences in phylogeographic patterns of flightless insect species occurring in different regions of Japan, we studied the phylogeography and demographic history of Silpha beetles occurring in cool-temperate habitats of two major islands, Honshu and Hokkaido, using sequences of the mitochondrial cytochrome oxidase subunit I (COI) gene. Honshu has a more mountainous topography, and cool-temperate habitats occur discontinuously, whereas Hokkaido, located to the north of Honshu, has more continuous cool-temperate habitats. A species endemic to Honshu, S. longicornis occurs on Honshu, whereas S. perforata occurs on Hokkaido and the East Asian continent. Our results indicate that the ancestors of S. longicornis colonized Honshu via a south-west route c . 0.7 Mya and the species has highly divergent populations in isolated mountainous areas of Honshu, whereas S. perforata colonized Hokkaido via a northern route less than 90 000 years ago and has less divergent geographic populations. During the last glacial period, S. perforata was probably restricted to refugia in southern Hokkaido and later expanded into northern Hokkaido, whereas S. longicornis populations existed in many isolated refugia, probably because of the complex topography of Honshu. Thus, our study demonstrates that, even between closely related species, interactions among biology, latitudinal climatic gradients and topography can produce different phylogeographic patterns.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 452–467.     

Composition,dynamics and disturbance regime of temperate deciduous forests in Monsoon Asia

Temperate deciduous forests in Monsoon Asia are classified into three forest types which differ in floristic composition, dynamics and disturbance regime. The cool temperate mixed deciduous broadleaf/conifer forest, dominated by Quercus spp. (mainly Q. mongolica or Q. crispula) and conifers, is distributed in northern parts of the temperate zone. The cool temperate deciduous forest, dominated by Fagus crenata, is distributed in Honshu, Japan under a humid climate through the year. The warm temperate deciduous forest dominated by Quercus spp. (mainly Q. acutissima or Q. serrata) occurs in the continental areas, the Korean Peninsula, and the Pacific Ocean side of Japan. The species diversity of cool temperate deciduous forest was lower than the other two types because of the intensive dominance of Fagus crenata. The disturbance regimes also varies among the three types; small scale treefall gaps are prevailing in the cool temperate deciduous forest, while larger scaled disturbances are important in the other two forest types. Fire seems to be important in the warm temperate deciduous forest, and both fire and large scale blowdowm seems important in the cool temperate mixed broadleaf/conifer forest. These differences in forest composition and disturbance regimes associated with climatic conditions and ancient human impacts have a close analogy with the Northeastern Hardwood forests in North America.     

The boreal forests of north-eastern Eurasia

The distribution, diversity and structure of boreal forests of monsoon Asia are reviewed. The boreal zone is circumscribed by the isotherms of 15°C and 45°C of Kira's Warmth Index (WI) and includes the territory of the Amur basin, the seashores of the Sea of Okhotsk, Kamchatka, Sakhalin and the Kuril islands. Boreal forests also occur on the mountains of Hokkaido, Honshu, Korea and NE China. The oceanic sectors of the boreal zone are composed mainly of birch forests, the marine sectors of spruce-fir forests and the continental sectors of larch and pine forests. The upper limit of the boreal forests ascends from the lowest level in the North to ca. 1500 m a.s.l. in the South of the zone, and reaches above 2500 m in the mountains of the temperate zone. In the boreal zone the secondary forests are common. They form after fires and cutting. Volcanism is an essential factor in the forest dynamics in the oceanic part of the region. It is suggested that as a result of global warming one may expect a strong alteration in vegetation pattern, especially in the northern and mountain areas.Paper presented at the International Symposium on Global Change Impacts on Terrestrial Ecosystems in Monsoon Asia (TEMA) Waseda University, Tokyo, 4–6 September 1993.     

Multiple origins of Locusta migratoria (Orthoptera: Acrididae) in the Japanese Archipelago and the presence of two major clades in the world: evidence from a molecular approach

Phylogenetic relationships among migratory locust (Locusta migratoria) populations in different climatic regions were analysed by sequencing four mitochondrial DNA regions, with special reference to the origin of Japanese populations. The populations are clearly separated into two clades: one consists of individuals from temperate and cold‐temperate areas of Japan and the Chinese continent, and the other comprises those from subtropical islands of Japan, Hainan Island in China, Timol Leste, Australia, Ethiopia, France, and some individuals from Tsushima Island and Honshu of Japan. The divergence time between the two clades is estimated to be 0.86–1.89 Mya. The phylogenetic analysis revealed that Japanese L. migratoria populations were composed of individuals of six different origins: (1) Hokkaido populations possibly from the Russian continent; (2) Honshu–Kyushu populations from the Chinese continent; (3) Southwest Island populations from Hainan Island or adjacent areas; (4) Ogasawara populations that might have originated from Micronesia; (5) part of the Tsushima population that originated from somewhere in the Asian tropics; and (6) a possible relict population of ancient southern haplotypes that exists in western areas of northern Honshu. The Tsugaru Straits and Tokara Straits have acted as effective geographical barriers, as in other organisms, isolating locust populations for a few thousand years. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 570–581.     

Late-quaternary shift offagus distribution

Based on six resolved maps of its pollen frequency (dating 13,000, 10,000, 8,000, 6,000, 3,000 and 500 yr B.P.) at 33 sites scattered almost throughout the Japanese Archipelago,Fagus shows principally a, unidirectional late-Quaternary shift in distribution from the southwest to the northeast. The maximum areal coverage was from 10,000 to 7,000 years ago, although northeastern populations were not fully developed by this time. The northern limit ofF. crenata, which has virtually not changed in the past 7,000 years, was probably regulated by total precipitation during the growing season from April to October (called the effective precipitation). ForF. crenata to migrate farther north in Hokkaido under the modern temperature regime, a minimum of 800 mm of precipitation, evenly distributed over this period, is required.     

Assessing the impact of land use and climate change on the evergreen broad-leaved species of <Emphasis Type="Italic">Quercus acuta</Emphasis> in Japan

To assess the impact of Quercus acuta, a dominant species in the evergreen broad-leaved forests of Japan, and its habitat shifts as a result of climate change, we predicted the potential habitats under the current climate and two climate change scenarios using a random forest (RF). The presence/absence records of Q. acuta were extracted from the Phytosociological Relevè Data Base as response variables, and four climatic variables (warmth index, WI; minimum temperature of the coldest month, TMC; summer precipitation, PRS; and winter precipitation, PRW) were used as predictor variables. The mean decrease in the Gini criterion revealed that WI was the most influential factor followed by TMC. The RF revealed a considerable increase in potential habitats (PHs) under the climate change scenarios for 2081–2100 (RCM20, 180,141 km²; MIROC, 175,635 km²) relative to the current climate (150,542 km²). The land use variables were used for masking PH. The PH masked by land use (PHLU) was approximately half of the PH under the current conditions (74,567 km²). Under the climate change scenarios and 1 km migration options, the PHLU were not increased relative to its value under the current conditions. The distribution of Q. acuta was restricted by the northward shift in northern Honshu, but expanded as a result of the upward shift into the mountain areas of Western Japan. Habitat fragmentation reduced the ability of migration to respond to climate change in the lowland areas of Japan.     

Contrasting patterns of nuclear microsatellite genetic structure of Fraxinus mandshurica var. japonica between northern and southern populations in Japan

Aim The aim of this study is to detect extant patterns of population genetic structure of Fraxinus mandshurica var. japonica in Japan, and to provide insights into the post‐glacial history of this species during the Holocene. Location Hokkaido and Honshu islands, Japan (including the Oshima and Shimokita peninsulas). Methods We examined nine polymorphic nuclear microsatellite loci to assess genetic variation within and among 15 populations across almost the entire range of the species in Japan. Extant patterns of geographical structure were analysed using Bayesian clustering, Monmonier’s algorithm, analysis of molecular variance, Mantel tests and principal coordinates analysis. Recent bottlenecks within populations and regional genetic variation were also assessed. Results Northern populations (Hokkaido Island and the Shimokita Peninsula) formed a single homogeneous deme, maintaining the highest level of allelic diversity on the Oshima Peninsula. By contrast, southern populations (Honshu Island) demonstrated strong substructure on both coasts. Specifically, populations on the Pacific side of Honshu exhibited significant bottlenecks and erosion of allelic diversity but preserved distinct subclusters diverging from widespread subclusters on the Japan Sea side of this island. Main conclusions Genetic evidence and life history traits suggest that F. mandshurica occupied cryptic northern refugia on the Oshima Peninsula during the Last Glacial Maximum, which is reflected in the species’ extant northern distribution. Strong geographical structure in southern populations, in agreement with fossil pollen records, suggests geographical isolation by mountain ranges running north–south along Honshu. Given that this tree species is cold‐adapted and found in riparian habitats, populations on the Pacific side of Honshu probably contracted into higher‐elevation swamps during warm post‐glacial periods, leading to a reduction of effective population sizes and rare allelic richness.     

The phylogeographic puzzle of Pseudoacanthocephalus toshimai,an amphibian acanthocephalan in northern Japan

The amphibian acanthocephalan, Pseudoacanthocephalus toshimai, was considered to be an island-endemic species in Hokkaido, Japan. However, the parasite was found from Rana ornativentris, Rana tagoi, Zhangixalus arboreus, and Bufo japonicus formosus in northern Honshu (Aomori and Iwate Prefectures), which is separated from Hokkaido by the Tsugaru Strait. The mitochondrial DNA-based phylogenetic and population genetic analyses of P. toshimai showed that the northern Honshu isolates are far distantly related to the Hokkaido isolates, and that a demographic population expansion occurred in Hokkaido during the recent geological past. The rich genetic diversity of P. toshimai in northern Honshu suggests a scenario that anuran hosts invaded Hokkaido together with P. toshimai via the land bridge of the Tsugaru Strait. However, the evolutionary history of Rana pirica, a main definitive host for P. toshimai in Hokkaido, is contradictory to the introduction scenario inferred from the parasite. The finding of several geographically mismatched isolates of P. toshimai from both northern Honshu and Hokkaido suggests a possibility that the migration of the parasite infrequently occurred between the two areas even after the land bridge disappeared. More detailed information on the evolutionary history of anurans is needed to resolve the biogeographical enigma of P. toshimai.     

Three species of the genus Ilyocryptus (Anomopoda, Branchiopoda) occurring in Japan

Three species of Ilyocryptus, Ilyocryptus sp., I. cuneatus Stifter 1988, and I. spinifer Herrick 1882, are newly recorded from the Japanese islands. The first two species were found separately at two sites in Hokkaido and the third at eight sites in Honshu and Kyushu. Their morphological features are briefly described based on parthenogenetic females. Received: May 21, 2001 / Accepted: July 23, 2001     

Allozyme Variation among Populations of Rana pirica (Amphibia: Anura)

Genetic variation of Rana pirica , an east Asian brown frog of the R. temporaria group, was elucidated by analysing 140 specimens from 11 populations from Hokkaido and Sakhalin, both locating near the eastern coasts of the Asian continent, and 12 specimens of R. ornativentris from Honshu, Japan mainland, as an outgroup, through horizontal starch gel electrophoresis. Rana pirica shows relatively low genetic differentiation, and high genetic affinities are found between Hokkaido and Sakhalin populations. Populations from these islands are morphologically somewhat differentiated but should be regarded as conspecific. Degree of genetic divergence among populations of amphibian species from Hokkaido, including R. pirica , is not so extensive as that reported for species from Japan mainland and relatively recent formation of amphibian fauna in Hokkaido is suggested.     

Genetic structure of Japanese populations of an ambrosia beetle,Xylosandrus germanus (Curculionidae: Scolytinae)

We examined the genetic structures of 13 Japanese populations of an ambrosia beetle, Xylosandrus germanus (Curculionidae: Scolytinae), to understand the effects of geographical barriers on the colonization dynamics of this species. The genetic structure was studied using portions of the mitochondrial cytochrome oxidase I (COI) gene. A phylogenetic analysis revealed three distinct lineages (clades A, B and C) within X. germanus. Clade A contained 21 haplotypes from all 13 populations; whereas clade B contained eight haplotypes from Hokkaido (Sapporo and Furano), Iwate and Nagano populations; and clade C contained only a single a haplotype from the Hokkaido (Furano) population. In the analysis of molecular variance (amova ), the greatest amount of genetic variation was detected between populations in Hokkaido and those in Honshu and other southern islands. Between these two groups of populations, all the values of the coefficient of gene differentiation were significantly larger than zero, except for the Hokkaido (Sapporo) versus Nagano comparison. Our results confirm that for X. germanus, gene flow has been interrupted between Hokkaido and Honshu since the last glacial maximum.     

Regulation of adult diapause inCoccinella septempunctata septempunctata andC. septempunctata brucki from two regions of Japan (A Minireview)

In similar climatic conditions of central/northern Europe and Hokkaido, Japan, both subspecies ofCoccinella septempunctata L. have similar life-cycles: long-day photoperiodic response ensures the induction of winter diapause. In the mild climate of Honshu, Japan,C. s. brucki Mulsant shows a different life cycle: short-day photoperiodic response leads to the induction of summer diapause while overwintering is quiescence. All populations show an important variation in photoperiodic response.     

Pseudoacanthocephalus toshimai sp. nov. (Palaeacanthocephala: Echinorhynchidae), a common acanthocephalan of anuran and urodelan amphibians in Hokkaido,Japan, with a finding of its intermediate host

The Ezo brown frog (Rana pirica) and the Ezo salamander (Hynobius retardatus) are endemic species of Hokkaido, the northernmost island of Japan. Intestinal adult acanthocephalans are common in these amphibians. A molecular identification based on nuclear and mitochondrial DNA markers demonstrated that the parasites from the anuran and the urodelan are the same species. In the neighboring Honshu island, another acanthocephalan from ranid frogs (e.g. Rana japonica and Rana ornativentris) has been identified as Acanthocephalus lucidus. The counterpart species from the amphibians of Hokkaido was morphologically indistinguishable from A. lucidus. However, clear genetic distinctiveness between the two allopatric populations (separated by islands) indicated the entity of a cryptic species. A phylogenetic tree inferred from sequences of 28S ribosomal DNA showed that the acanthocephalans from Honshu and Hokkaido belong to the genus Pseudoacanthocephalus. Therefore, Pseudoacanthocephalus toshimai sp. nov. is proposed for the cryptic species in Hokkaido, together with the transfer of A. lucidus in Honshu to Pseudoacanthocephalus lucidus comb. nov. The present field survey further demonstrated Ligidium japonicum, an isopod crustacean living in the litter layer of forests, to be an intermediate host of the new species.     

Ground crickets singing in volcanic warm “islets” in snowy winter: Their seasonal life cycles,photoperiodic responses and origin

The ground cricket Dianemobius nigrofasciatus overwinters as an egg in Japan, being univoltine in Hokkaido and northern Honshu and bivoltine farther south. In Hokkaido, however, this cricket is heard singing in winter in several fumarolic fields covered with moss and grasses locally known as “bokke”. In such warm “islets” the adult density was high in early summer and again in autumn, indicating that the cricket is bivoltine in contrast to the univoltine life cycle outside the bokke habitats in Hokkaido. Eggs laid by females collected at regular intervals from a bokke habitat showed a clear seasonal cycle of diapause incidence. At 26°C, the bokke strains produced non‐diapause eggs under long days and diapause eggs under short days as in the southern bivoltine populations, although the critical day‐length was longer than in the south. Several strains derived from non‐bokke habitats in Hokkaido and northern Honshu produced high percentages of diapause eggs under long days as well as short days as expected for the univoltine life cycle. Winter adults singing in bokke habitats could be either survivors of the autumn generation or individuals derived from eggs laid in autumn and then matured in response to the high soil temperature. In the laboratory, the proportion of egg diapause in short days was decreased by selection only for several generations. Phylogenetic trees of bokke and non‐bokke populations, based on both the nucleotide sequence of the mitochondrial COI gene and four allozyme loci, suggest that bokke populations have not been isolated from non‐bokke populations for an evolutionarily significant time.     

Two Genetically Distinct Lineages of the Sika Deer, Cervus nippon, in Japanese Islands: Comparison of Mitochondrial D-Loop Region Sequences

To investigate genetic diversity among populations of the sika deer, Cervus nippon, nucleotide sequences (705–824 bases) of the mitochondrial D-loop regions were determined in animals from 13 localities in the Japanese islands. Phylogenetic trees constructed by the sequences indicated that the Japanese sika deer is separated into two distinct lineages: the northern Japan group (the Hokkaido island and most of the Honshu mainland) and the southern Japan group (a part of the southern Honshu mainland, the Kyushu island, and small islands around the Kyushu island). All sika deer examined in this study shared four to seven units of repetitive sequences (37 to 40 bases each) within the D-loop sequences. The number of tandem repeats was different among the populations, and it was specific to each population. Six or seven repeats occurred in populations of the northern Japan group, while four or five repeats occurred in populations of the southern Japan group. Each repeat unit included several nucleotide substitutions, compared with others, and 26 types were identified from 31 animals. Sequences of the first, second, and third units in arrays were clearly different between the northern and the southern groups. Based on these D-loop data, colonization and separation of the sika deer populations in the Japanese islands were estimated to have occurred less than 0.5 million years before present. Our results provide an invaluable insight into better understanding the evolutionary history, phylogeny, taxonomy, and population genetics of the sika deer.     

The genetic structure of <Emphasis Type="Italic">Quercus crispula</Emphasis> in northeastern Japan as revealed by nuclear simple sequence repeat loci

Previous studies have reached different discussions about the genetic variation and genetic structure of Quercus crispula populations in northeastern Japan. This is a common oak species in Eastern Asia. Some studies have suggested that the populations in northeastern Japan were derived from those remaining in the southwest after the last glacial maximum (LGM), whilst other studies have found evidence that populations persisted in northeastern Japan during the LGM. Using seven highly polymorphic nuclear simple sequence repeat loci, we investigated the genetic structure of 16 Q. crispula populations along a latitudinal gradient in northeastern Japan (northern Honshu and Hokkaido), spanning about half of the species’ biogeographic range in the country. Although the level of population differentiation was low (F _ST = 0.021; G^￠_\textST G^{\prime}_{\text{ST}}  = 0.090), two geographically differentiated clusters were detected by STRUCTURE analysis. The first cluster included most of the populations in Hokkaido, and may indicate continued survival throughout past glacial periods. We found a significant decrease in allelic richness with latitude, so the second cluster may represent an expansion of the lineage from Honshu during the post-glacial period. These results should enhance our understanding of historical north–south migrations of this species in northeastern Japan.     

Further Analysis of Intraspecific Sequence Variation of Chloroplast DNA in Primula cuneifolia Ledeb. (Primulaceae): Implications for Biogeography of the Japanese Alpine Flora

Primula cuneifolia Ledeb. (Primulaceae), we analyzed intraspecific variation of the nucleotide sequences of non-coding regions of chloroplast DNA: the intergenic spacers between trnT (UGU) and the trnL (UAA) 5′exon, the trnL (UAA) 3′exon and trnF (GAA), and atpB and rbcL. In 20 populations of P. cuneifolia, 22 nucleotide substitutions and five insertions/deletions were inferred, and their genetic distances ranged from 0.001 to 0.008. Eight distinct haplotypes could be recognized and each haplotype was found to be geographically structured. Three major clades (the Northern, Hokkaido and Southern clades) were revealed in phylogenetic analyses of the haplotypes. The haplotypes of the Northern clade had a wider distribution area in the populations of Mt. Rausu and Rishiri Island of eastern and northern Hokkaido in Japan, northward to Unalaska Island in the Aleutians, and those of the Hokkaido clade were distributed in the populations of central Hokkaido and Mt. Iwaki of the northern Honshu in Japan; in addition, those of the Southern clade were observed only in the populations of the central Honshu. It was shown that the genetic diversifications of the Southern clade were higher than those of the Northern and Hokkaido clades. Furthermore, it was shown that the topology within the Southern clade was hierarchical, and the haplotypes of the Southern populations in the clade were derivative. From these results, we concluded that the cpDNA haplotypes of the three clades in P. cuneifolia arose and assumed the present distribution areas through several cycles of glacial advance and retreat in the Pleistocene. Received 24 June 1998/ Accepted in revised form 28 December 1998