A low <Emphasis Type="Italic">Platypus quercivorus</Emphasis> hole density does not necessarily indicate a small flying population

The ambrosia beetle Platypus quercivorus causes mass mortality of Fagaceae trees in Japan, and tree species differ in their susceptibility to P. quercivorus. We hypothesized that interspecific differences in susceptibility are caused by differences in beetle infestation patterns, that is, how many beetles fly to a tree and how many of those arriving bore holes. To examine how tree susceptibility is related to these parameters, two tree species with different degrees of susceptibility were studied (highly susceptible Quercus crispula and less susceptible Q. salicina). Specifically, we measured the number of male beetles per unit area that flew to the host trees (NFM) and the density of holes bored by male beetles (DH). From these two values, we calculated the proportion of male beetles flying to a host tree that bored holes (PBM). These parameters were compared for the two Quercus species. Although the two species did not differ in NFM in 2003, PBM was markedly lower for Q. salicina than for Q. crispula and DH was significantly lower for Q. salicina than for Q. crispula. Thus, the lower susceptibility of Q. salicina is partly explained by its low PBM.     

Types of frass produced by the ambrosia beetle <Emphasis Type="Italic">Platypus quercivorus</Emphasis> during gallery construction,and host suitability of five tree species for the beetle

We studied the biology of the ambrosia beetle Platypus quercivorus in the logs of five tree species to determine: (1) the relationship between the amount of frass produced by beetles and tunnel length, (2) the relationship between frass shape and the morphological characteristics of P. quercivorus mouthparts, and (3) the suitability of five tree species for P. quercivorus reproduction. Five logs each from healthy Quercus crispula, Q. serrata, Castanea crenata, Sorbus japonica, and Cryptomeria japonica trees were used in this experiment. The results showed that there was a linear relationship between the amount of frass and tunnel length. Whenever powdery frass was produced, larvae were found in the gallery in the log, while fibrous frass was present only in galleries that contained just adults. The mouthparts of adults were completely sclerotized, which likely accounts for the fibrous frass production. Host preference of P. quercivorus was examined at two stages using five tree species. The first stage is digging initiation, which concerns male preference for digging a tunnel. More holes were made by males on S. japonica and Q. serrata logs, while fewer holes were made on C. japonica logs. The second stage is characterized by female orientation, mating, and progeny development. Platypus quercivorus could complete its lifecycle only in the two Quercus spp.     

Aggregation ofPlatypus quercivorus (Murayama) (Coleoptera: Platypodidae) on oak logs bored by males of the species

To provide evidence for the aggregation ofPlatypus quercivorus (Murayama) (Coleoptera: Platypodidae) in association with the mass mortality of oak trees in Japan, we compared the number of beetles attracted to oak logs bored by males with the number of beetles attracted to logs that had not been bored. Large numbers of males and females were attracted to the bored logs, and the total numbers increased as the number of entry holes in the logs increased. This result shows that both male and female beetles aggregate on logs bored by the males. We collected a large number of beetles attracted to living trees bored by the beetles, and the total number of beetles collected increased as the number of entry holes/m² on the tree trunk increased. Beetles also aggregated on living trees bored by the males.     

Effects of previous forest types and site conditions on species composition and abundance of naturally regenerated trees in young <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> plantations in northern Japan

Naturally regenerated trees in young Cryptomeria japonica plantations were investigated in 141 quadrats of 10 × 50 m within a watershed of 1,000 ha, and factors affecting their composition and abundance were examined. The species composition of naturally regenerated trees was classified into four types. Dominant species were Swida controversa, Magnolia hypoleuca and Pterocarya rhoifolia in Type A1, Betula maximowicziana, M. hypoleuca, Quercus crispula and Castanea crenata in Type A2, Q. serrata and C. crenata in Type B1, and Pinus densiflora in Type B2. The results of path analysis showed significant influences of previous forest type and geology among the factors that correlated with the species composition of naturally regenerated trees; previous forest type in particular showed a higher absolute path coefficient value. Species composition types of naturally regenerated trees corresponded to the previous forest types: Types (A1 and A2) corresponded to the natural forests composed of Thujopsis dolabrata var. hondai, Fagus crenata, Aesculus turbinata, etc., Type B1 corresponded to the secondary Q. crispula and Q. serrata forest, and Type B2 corresponded to the secondary P. densiflora forest, respectively. The abundance of naturally regenerated trees was strongly affected by geology; i.e., plantations on soft-sedimentary dacitic tuff contained great amounts of colonizing P. densiflora trees.     

Predicting the spatial distribution of major species composition in secondary hardwood forests on Mt. Gozu,central Japan,based on environmental factors

Our goal was to predict the spatial distribution of canopy species composition in secondary deciduous hardwood forests at a fine spatial resolution, based on climatic and topographical factors using a geographic information system. We studied secondary forests on Mt. Gozu, Niigata Prefecture, central Japan. Canopy species composition was investigated in 100 sample plots within the study area. A digital elevation model (DEM) was created, and topographical, hydrological, and light factors were calculated using the DEM. Climatic factors were interpolated by kriging. The five major species used as response variables were Fagus crenata, Quercus serrata, Quercus crispula, Magnolia obovata, and Castanea crenata. We prepared three possible explanatory variable sets: climatic variables only, both climatic and topographic variables, and topographic variables only. Multivariate regression trees were derived, and the accuracy of predicting the major species composition was tested. The multivariate regression trees derived from the climatic variable set and from the climatic and topographic variable set had better accuracy than the regression tree derived from the topographic variable set. In the regression tree generated by the climatic and topographic variable set, the warmth index was the principal explanatory variable in classifying forest types, followed by topographic factors. This regression tree would be preferable to the other two regression trees for the prediction of canopy species composition.     

Differences in leafminer (Phyllonorycter, Gracillariidae, Lepidoptera) and aphid (Tuberculatus, Aphididae, Hemiptera) composition among Quercusdentata, Q. crispula, Q. serrata, and their hybrids

Leafminer (Phyllonorycter, Gracillariidae, Lepidoptera) and aphid (Tuberculatus, Aphididae, Hemiptera) composition were studied in three deciduous oak species, Quercus dentata, Q. crispula, and Q. serrata, and their hybrids in Tomakomai Experimental Forest of Hokkaido University, Hokkaido, northern Japan. Identification of trees in this forest was done mainly on the basis of discriminant analysis on leaf morphology with reference to trees in pure Q. dentata and Q. crispula stands and a Q. serrata stand mixed with Q. crispula. The results suggested that hybridization occurred in all combinations (i.e. Q. dentata–Q. crispula, Q. crispula–Q. serrata, and Q. serrata–Q. dentata) and the frequency of hybrids was approximately 10%. The composition of Phyllonorycter and Tuberculatus species differed between Q. dentata and Q. crispula or Q. serrata, but did not differ between Q. crispula and Q. serrata. Thus, Q. dentata could differ from Q. crispula and Q. serrata in chemical properties that determine herbivore host selection, survival, and performance, possibly reflecting eco-physiological differences or phylogenetic distances. The study insects were divided into three groups: species specialized to Q. dentata (three Phyllonorycter and one Tuberculatus species), those to Q. crispula and Q. serrata (six Phyllonorycter and two Tuberculatus species), and a species collected at least from Q. dentata and Q. crispula (one Tuberculatus species). Putative hybrid trees of Q. dentata and Q. crispula harbored both Q. dentata-specific and Q. crispula-specific insects.     

Changes in water status in seedlings of six species in the Fagaceae after inoculation with Raffaelea quercivora Kubono et Shin-Ito

To clarify the differences in susceptibility of six species in the Fagaceae (Quercus crispula, Quercus serrata, Quercus acutissima, Quercus phillyraeoides, Quercus glauca, and Castanopsis cuspidata var. sieboldii) to Raffaelea quercivora, we inoculated this fungus on seven potted 5-year-old seedlings of each species, observed symptom development, and measured xylem pressure potential (XPP) after inoculation. The first death was observed on the 11th day in Q. crispula and on the 56th day in Q. serrata. The number of dead seedlings of Q. crispula and Q. serrata were five and one, respectively, whereas no mortality was observed in the other four species. The XPP of inoculated seedlings in both Q. crispula and Q. serrata decreased after inoculation. In contrast, the XPP of inoculated seedlings of the other four species remained almost the same as in the control seedlings. These results indicate that R. quercivora is pathogenic to Q. crispula and Q. serrata and that the susceptibility of the six Fagaceae species in our study differed among species.     

Genotype distribution of Raffaelea quercivora in the oak galleries and its composition in the mycangia of Platypus quercivorus

Raffaelea quercivora is the pathogenic fungus that causes Japanese oak wilt. The female monogynous ambrosia beetle, Platypus quercivorus, carries this fungus in mycangia on the pronotum. These beetles bore galleries in oak trees with their partners to produce offspring, and they deposit fungus on the gallery walls from their mycangia. The offspring mature in the gallery, before loading the fungal pathogen and flying from the gallery to other healthy trees. To investigate the unloading and loading modes of the fungus within the gallery, we developed four polymorphic microsatellite markers for R. quercivora and identified the fungal genotypes in the galleries and mycangia of the beetles. Small wood chips were sampled at 5–10‐mm intervals from the walls of five galleries in a dead Quercus serrata tree. The pronota were also sampled from five female adult beetles. The genotypes of the R. quercivora isolates from the wood chips and pronota were identified using the microsatellite makers. The genotypic analysis showed that each gallery was inhabited patchily by 5–10 genotypes of R. quercivora, and the mycangia of each beetle contained 3–6 genotypes. These results indicate that diverse R. quercivora genotypes are unloaded repeatedly from the mycangia of female beetles onto the gallery wall, which results in their patchy distribution on the walls. When the offspring leave the host tree, the fungal clones that proliferate in the walls are also loaded repeatedly into the mycangia of the mature beetles.     

Spatial distribution pattern of attack of the oak borer,Platypus quercivorus (Murayama) (Coleoptera: Platypodidae), and scolytid ambrosia beetles (Coleoptera: Scolytidae) on fresh logs

The spatial distribution patterns of the attack on fresh logs ofPasania edulis was studied for the oak borer,Platypus quercivorus (Murayama), and two species of Scolytid ambrosia beetles,Xylosandrus crassiusculus (Motschulsky) andXyleborus attenuatus Blanford, in 1994 and 1995. On the logs where onlyP. quercivorus attacked, the entry holes were distributed uniformly when attack intensity was low. However, the distribution pattern became more aggregated with the increase in attack intensity. On logs where bothP. quercivorus and the two Scolytid species attacked, there was a negative association between the spatial distribution of the entry holes ofP. quercivorus and that of scolytids. Simultaneous attack of two scolytids also increased the degree of aggregation of the entry holes ofP. quercivorus. The entry holes of scolytids were distributed in groups irrespective of the attack intensity of scolytids andP. quercivorus. These results suggest an asymmetrical interspecific relationship betweenP. quercivorus and scolytids. Concentration of the entry holes ofP. quercivorus in a small area may cause a considerable decline in the reproductive success in the galleries constructed there.     

Discoloured and non‐conductive sapwood among six Fagaceae species inoculated with Raffaelea quercivora

Mass mortality of oak trees has been occurring in Japan since the late 1980s. The fungus Raffaelea quercivora has been frequently isolated from discoloured sapwood in dead or wilting trees and inoculation experiments have shown it to be capable of causing wilting and xylem discoloration in several oak species, notably Quercus crispula and Q. serrata. In this study, we inoculated seedlings of six Fagaceae species with R. quercivora and, after 56 days, measured the vertical length of the discoloration and the areas of discoloured and non‐conducting sapwood on stem cross‐sections. The sapwood discoloration and the water non‐conduction areas were larger in Q. crispula and Q. serrata than in the other species.     

Horizontal niche differentiation of ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) within the standing trunk of <Emphasis Type="Italic">Quercus serrata</Emphasis>

Niche differentiation, in terms of time and space, has been reported within a community of ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) in a tree trunk. Two platypodid ambrosia beetles, Platypus quercivorus (Murray) and Platypus calamus Blandford, utilize a similar height range with respect to the Quercus serrata Murray (Fagaceae) trunk. The sapwood and heartwood (i.e., wood types) are known to differ in their physical and chemical characteristics. The objective of this study was to determine the differences among ambrosia beetles, in terms of wood type preferences, in a Q. serrata tree trunk. We analyzed the horizontal distribution patterns of ambrosia beetles within two mature Q. serrata tree trunks. Platypus calamus and the other ambrosia beetles [P. quercivorus, Ambrosiodmus lewisi (Blandford), Ambrosiophilus atratus Eichhoff, and Xyleborus sp.] showed different horizontal distribution patterns within the wood. The former tended to utilize the heartwood, whereas the latter utilized the sapwood. These results suggest that the wood type can be regarded as one of the niche dimensions of ambrosia beetles in a Q. serrata trunk.     

Responses ofQuercus sapwood to infection with the pathogenic fungus of a new wilt disease vectored by the ambrosia beetlePlatypus quercivorus

Quercus serrata andQ. crispula wilt during the summer in wide areas along the Sea of Japan. Mass attacks of trees by an ambrosia beetle (Platypus quercivorus) are characteristic before appearance of the wilting symptoms. This study investigated the pathogenic effects of a fungus detected specifically in the wilting trees. This hyphomycete fungus,Raffaelea sp., has a distribution that correlates with the discolored xylem area called wound heartwood in which vessels are dysfunctional. Tylosis formation around the hyphae indicates vessel dysfunction. In areas with discoloration, the fungal hyphae were invading living ray parenchyma cells from the vessel lumen. As a protective reaction the ray cells exuded yellow substances into the vessels, but these substances seemed ineffective against the fungal activity, probably because the fungus disperses along the beetle's gallery before enough substance can accumulate. It should allow wide discoloration in sapwood. Cambium was not necrotic around the fungus. The cytological process in the host was as follows: (1) synthesis of secondary metabolites by the stimuli of oak fungus; (2) exudation of yellow substances into vessels; and (3) dysfunction of vessels and wound heartwood formation. In regard to wilting of trees, the pathogenicity of the fungus should be assessed by its ability to stop sap flow.     

Effects of crown release on basal area growth rates of some broad-leaved tree species with different shade-tolerance

We compare three or four years effects of crown release on stem growth rates among four deciduous broad-leaved tree species,Fagus crenata, Quercus crispula, Magnolia obovata andAcer mono, in an even-aged coppice forest, central Japan. The crown release significantly improved relative growth rates for basal-area (RGR_BA) of the four species. However the effects of neighboring gap area (GAPA) on the RGR_BA differed among the species. The effects were significant forQ. crispula andM. obovata, indicating that growth rates of these two species increase with intensity of the crown release. In contrast, the RGR_BA ofF. crenata andA. mono were not correlated with the GAPA, suggesting that their growth rates are unrelated to the intensity. We considered that such differences among species were closely related to their photosynthetic characteristics of light use; less-tolerant species (Q. crispula andM. obovata) exhibit greater growth rates relative to the intensity of crown release than tolerant species (F. crenata andA. mono). Based on these results, we proposed implications for thinning practices in mixed forests of species with different shade-tolerance.     

Relationships between mass mortality of two oak species (Quercus mongolica Turcz. var.grosseserrata Rehd. et Wils. andQ. serrata Thunb.) and infestation by and reproduction ofPlatypus quercivorus (Murayama) (coleoptera: Platypodidae)

This study investigated the number of adults ofPlatypus quercivorus (Murayama) captured on host oak (Quercus spp.) trees, the attack density (the number of entry holes per 100 cm²), and the number of emerging adults to clarify the relationships between the beetle’s colonization on trees in oak stands and tree mortality. The initial attack ofP. quercivorus, which was the most intense attack, was observed on most living trees simultaneously. Although some attacked trees died within the year of the attack or in the next year, a high density of attack did not usually cause the death of host trees. Surviving trees suffered low levels of consecutive attack after the initial attack. BecauseP. quercivorus successfully produced broods only in the dead trees, the population of the insect seems to be maintained only in stands where oak mortality occurs. However, the adults that landed on most of the surviving trees appeared unable to reproduce probably due to degradation of host quality. Thus, oak mortality probably ceases within 3 or 4 years after the start of infestation in a stand, with subsequent reductions in population density of the borer.     

Comparisons of hyphal growth of Raffaelea quercivora among four Japanese Fagaceae species

Japanese oak wilt causes widespread oak mortality in Japan. Possible differences in susceptibility to the causal fungus, Raffaelea quercivora, may be due to vessel arrangements in host trees. To clarify whether constitutive defence mechanisms including vessel arrangements or induced defence mechanisms are the main determinants of host susceptibility, we inoculated the fungus into living seedlings or sterilized stem segments of four Japanese fagaceous species. In seedlings, water conductance was assessed with dye. In both seedlings and stem segments, hyphal growth was examined by fluorescence microscopy. In seedlings, the area of non‐conductive sapwood in stem cross sections and hyphal growth differed significantly among species. In the susceptible species Quercus crispula and Quercus serrata, hyphal growth was significantly and positively correlated with the proportion of non‐conductive sapwood. In stem segments, hyphal growth was not significantly different among species or between vessel arrangements and was similar to or greater than that in seedlings. These results suggest that the extent of sapwood colonization by R. quercivora could be used as a marker for susceptibility and that susceptibility is determined mainly by induced defence responses.     

Individual-scale responses of five dominant tree species to single-tree selection harvesting in a mixed forest in Hokkaido, northern Japan

We investigated the individual-scale responses of five dominant species (Abies sachalinensis, Acer mono, Tilia japonica, Quercus crispula, and Betula ermanii) to single-tree selection harvesting in a conifer–broadleaved mixed forest in Hokkaido, northern Japan. Using data from stems with a diameter at breast height of ≥12.5 cm, collected during 20 years of monitoring a 6.7-ha stand, we analyzed the effects of harvesting in the neighborhood on tree recruitment and the growth (diameter class transition) and mortality of the residual trees. The effects of harvesting varied considerably among tree sizes and species. Harvesting improved the recruitment of A. mono and B. ermanii, and moderated the negative effect of the initial basal area of the surrounding canopy trees on the recruitment of Q. crispula. Conversely, harvesting limited the recruitment of A. sachalinensis by offsetting the positive effect of the initial basal area of the surrounding canopy trees. The growth of A. sachalinensis and Q. crispula decreased with the initial basal area of the surrounding canopy trees. Harvesting in the neighborhood resulted in an improvement in the growth of the trees of these species only in the smaller size classes. With increasing local harvesting intensity, the mortality of smaller A. sachalinensis trees decreased, whereas the mortality of larger trees increased. These results suggest that differences in the local harvesting intensity, spatial patterns of harvesting, and initial stand structures influence the stand-scale dynamics in response to partial harvesting in the mixed forests of this region.     

The stand dynamics of a mixed coppice forest of shade-tolerant and intermediate species

We investigated stand dynamics of a mixed coppice forest of shade-tolerant and intermediate species, Fagus crenata Bl. and Quercus crispula Bl., in five stands with canopy-stem densities from 420 to 1320 ha⁻¹ (estimated stand age: 37–62 years). The mean basal area (BA), exposed-crown area (ECA) and crown depth (CD) per stem were significantly larger in F. crenata in the three lower density plots. Also Q. crispula had considerably less overtopped stems in the two lower density plots. The mean annual BA increments of the two species were similar in the earlier 20 years; however, in the most recent 5 years, F. crenata had a higher relative growth rate for basal area (RGR_BA) than Q. crispula. The most critical variable of the RGR_BA was the ECA for F. crenata and the relative height (RH) for Q. crispula. We predict that dominance of F. crenata will gradually increase over that of Q. crispula as stands mature. This seems to be caused by lower shade-tolerance despite the similar initial growth rates of Q. crispula compared with F. crenata.     

Virulence of Raffaelea quercivora isolates inoculated into Quercus serrata logs and Q. crispula saplings

This study was conducted to compare the virulences of various isolates of Raffaelea quercivora—a fungus that causes Japanese oak wilt disease—towards Japanese oak trees. Five isolates were collected from a wide range of Japan and inoculated into Quercus serrata logs and Q. crispula saplings. The tangential length of the discolored sapwood in the Q. serrata logs differed significantly among the isolates. The trend in isolate virulence was similar for the Q. serrata logs and the Q. crispula saplings. This is the first report suggesting that there is variability in virulence among isolates of R. quercivora.     

Manipulation of tannins in oaks by galling cynipids

We measured the protein precipitation capacity of tannins in the tissues of Quercus crispula and Quercus serrata galls induced by a cynipid (Trigonaspis sp.) to examine the nutrition hypothesis on gall induction. The protein precipitation capacity was significantly lower in the nutritive tissues of galls, on which the cynipids feed, than in ungalled sound leaves and in the outer gall tissues. In addition, the protein precipitation capacities in the leaves and in the outer gall tissues were significantly larger in Q. crispula than in Q. serrata samples, whereas that of the nutritive tissues did not differ between these oak species. These results suggest that gall induction is one of the adaptations of galling cynipids to host–plant defensive compounds, and support the nutrition hypothesis.     

Effects of mating on the termination of aggregation by the oak borer,Platypus quercivorus (Murayama) (Curculionidae: Platypodinae) during host colonization

We conducted a field experiment to determine whether logs bored by male Platypus quercivorus beetle were less attractive to conspecifics after beetles had mated. Bored (with male beetles) and unbored logs were placed in a beetle-infested forest and enclosed within a cage. For half of the bored logs, female beetles were released to mate with the male beetles. Log attractiveness was determined by assessing the number of beetles captured on adhesive paper placed on each cage enclosing the log. On the logs on which female beetles were released, the number of beetles captured decreased distinctly after release, and did not differ from the number captured on the unbored logs. In contrast, the number of beetles captured on logs with only unpaired males was higher than the number captured on the unbored logs. This result confirms that the attractiveness of logs bored by male P. quercivorus beetles declines after mating occurs. To our knowledge, this is the first evidence for the relaxation of aggregation pheromone following mating for beetles in the subfamily Platypodinae.