Raffaelea quercivora sp. nov. associated with mass mortality of Japanese oak, and the ambrosia beetle (Platypus quercivorus)

 A hyphomycete consistently isolated from dead oak trees (Quercus serrata and Q. mongolica var. grosseserrata) attacked by the ambrosia beetle Platypus quercivorus in Japan is described and illustrated as Raffaelea quercivora sp. nov. The new species is characterized by having small obovoid to pyriform sympodioconidia and slender, long conidiophores that taper to a point. The fungus has been isolated from the body surfaces and mycangia of the beetle. It is likely that the fungus was transferred to oak trees by P. quercivorus. Received: August 20, 2001 / Accepted: March 14, 2002     

<Emphasis Type="Italic">Candida kashinagacola</Emphasis> sp. nov., <Emphasis Type="Italic">C. pseudovanderkliftii</Emphasis> sp. nov. and <Emphasis Type="Italic">C. vanderkliftii</Emphasis> sp. nov., three new yeasts from ambrosia beetle-associated sources

Three new yeast species, Candida kashinagacola (JCM 15019(T) = CBS 10903(T)), C. pseudovanderkliftii (JCM 15025(T) = CBS 10904(T)), and C. vanderkliftii (JCM 15029(T) = CBS 10905(T)) are described on the basis of comparison of nucleotide sequences of large subunit ribosomal DNA D1/D2 region (LSU rDNA D1/D2). The nearest assigned species of the three new species was Candida llanquihuensis. Candida kashinagacola and C. pseudovanderkliftii differed from C. llanquihuensis by 3.8% nucleotide substitution of the region, while C. vanderkliftii did by 4.4%. Three new species differed in a number of physiological and growth characteristics from any previously assigned species and from one another. A phylogenetic tree based on the sequences of LSU rDNA D1/D2 showed that these new species together with Candida sp. ST-246, Candida sp. JW01-7-11-1-4-y2, Candida sp. BG02-7-20-001A-2-1 and C. llanquihuensis form a clade near Ambrosiozyma species. The new species did not assimilate methanol as a sole source of carbon, which supported the monophyly of these non methanol-assimilating species which are closely related to the methylotrophic yeasts. Candida kashinagacola was frequently isolated from the beetle galleries of Platypus quercivorus in three different host trees (Quercus serrata, Q. laurifolia and Castanopsis cuspidata) located in the sourthern part of Kyoto, Japan, thus indicating that this species may be a primary ambrosia fungus of P. quercivorus. On the other hand, C. pseudovanderkliftii and C. vanderkliftii were isolated only from beetle galleries in Q. laurifolia. Candida vanderkliftii was isolated from beetle gallery of Platypus lewisi as well as those of P. quercivorus. Candida pseudovanderkliftii and C. vanderkliftii are assumed to be auxiliary ambrosia fungi of P. quercivorus.     

Patterns of community composition and diversity in latent fungi of living Quercus serrata trunks across a range of oak wilt prevalence and climate variables in Japan

Given that forest dieback due to emerging pests is increasing under global warming, understanding the relationships between pests, climate, and forest biodiversity is an urgent priority. In Japan, mass attacks of an ambrosia beetle, vectoring a pathogenic fungus, cause oak wilt outbreaks in recent decades. Here, the associations of oak wilt and climate with wood-inhabiting fungal communities in apparently healthy Quercus serrata trunks were investigated using DNA metabarcoding in seven sites along a climatic gradient in Japan. Amplicon sequencing of the fungal internal transcribed spacer 1 region generated 1,339,958 sequence reads containing 879 fungal operational taxonomic units (OTUs) in 234 wood samples. OTU compositions were significantly different between sites with and without oak wilt. OTU richness increased with temperature and precipitation at sites where oak wilt was present, but this relationship was not observed at sites without oak wilt, possibly due to interaction between oak wilt and climate.     

Attraction of ambrosia and bark beetles to coast live oaks infected by Phytophthora ramorum

1 Sudden oak death is caused by the apparently introduced oomycete, Phytophthora ramorum. We investigated the role of bark and ambrosia beetles in disease progression in coast live oaks Quercus agrifolia. 2 In two Marin County, California sites, 80 trees were inoculated in July 2002 with P. ramorum and 40 were wounded without inoculation. Half of the trees in each group were sprayed with the insecticide permethrin [cyclopropanecarboxylic acid, 3‐(2,2‐dichloroethenyl)‐2,2‐dimethyl‐(3‐phenoxyphenyl) methyl ester] to prevent ambrosia and bark beetle attacks, and then were sprayed twice per year thereafter. After each treatment, sticky traps were placed on only the permethrin‐treated trees. Beetles were collected periodically in 2003. 3 Inoculated trees accounted for 95% of all beetles trapped. The ambrosia beetles Monarthrum scutellare and Xyleborinus saxeseni and the western oak bark beetle Pseudopityophthorus pubipennis were the most abundant of the seven species trapped. 4 Permethrin treatment delayed initiation of beetle attacks and significantly reduced the mean number of attacks per tree. Beetles did not attack any wounded or noncankered inoculated trees. 5 Trees with larger cankers trapped more beetles early in the disease. Once permethrin lost effectiveness, the number of beetle entrance tunnels was a more reliable predictor of subsequent trap catch than was canker size. 6 Beetles were initially attracted to P. ramorum cankers in response to kairomones generated in the host‐pathogen interaction. After beetles attacked the permethrin‐treated trees, aggregation pheromones most probably were the principal factor in beetle colonization behaviour.     

Guild structure of ambrosia beetles attacking a deciduous oak tree Quercus serrata in relation to wood oldness and seasonality in three locations in the Central Japan

In order to determine factors influencing ambrosia beetle guilds on Quercus serrata, we investigated ambrosia beetles guilds by using Q. serrata bait logs in three locations in the Central Japan. Timing of cutting trees and timing of exposure were artificially controlled. Influences of location, timing of cutting, timing of exposure and wood oldness on species richness, abundances and guild structure were analyzed. Species richness and abundance peaked on bolts prepared in April–May, on bolts exposed in July, and on 2–3‐month‐old bolts. Eliminating greatest influences of location on abundance, results of hierarchical partitioning showed that timing of cutting trees had a strong influence on both species richness and abundance. LOC‐A (Aichi), in which Japanese oak wilt disease incidence occurred, showed the greatest species richness and the smallest value of Pielou's evenness. Abundance of the most major species was more than twice that of the second major species, which was a likely cause of the smallest evenness in LOC‐A. Trees killed by the Japanese oak wilt disease may have increased the abundance of the major species. On the contrary, in LOC‐C (Chichibu), alpha and beta diversity both given by Shannon index and Pielou's evenness were greatest among the three locations although species richness was smallest. High similarity between guilds in LOC‐A and LOC‐B (Chiba) was probably caused by similarity in vegetation. The location had the greatest effect on determining guild structure. Effect of timing of exposure was greater than timing of cutting. The effect of wood oldness was negligible. A hierarchical structure among the three factors was a likely cause of their relative importance determining guild structure.     

Microbial symbiotes of the ambrosia beetleXyloterinus politus

Progression in the understanding of the microecology of ambrosia beetles and their associated microorganisms is briefly reviewed. Between the 1840s and the early 1960s the concept of one ambrosial fungus per ambrosia beetle was emphasized. Some subsequent research has supported the view that each ambrosia beetle plus several associated microorganisms constitute a highly co-evolved symbiotic community. It was hypothesized in this study that such a community of symbiotic microbial species, not just one ambrosial fungus, is actively cultivated and perpetuated by the ambrosia beetleXyloterinus politus. Experimental results indicated that bacteria, yeasts, a yeastlike fungus, and ambrosial fungi compose such a symbiotic microbial complex in association withX. politus. The microecology of the ectosymbiotic microorganisms in relation to this insect is discussed.     

Platypus quercivorus ambrosia beetles use leaf volatiles in host selection

Although chemical volatiles emitted from host and non-host trees have been suggested as important cues for bark and ambrosia beetles, their responses to leaf volatiles is poorly understood. The oak ambrosia beetle, Platypus quercivorus (Murayama) (Coleoptera: Curculionidae), is a vector for the fungus that causes Japanese oak wilt. Using a Y-tube olfactometer, we tested the behavioral response of P. quercivorus to leaf volatiles emitted from four host trees – Quercus crispula Blume, Quercus serrata Murray, Quercus salicina Blume, and Castanea crenata Sieb. & Zucc. (all Fagaceae) – and two non-host trees, Fagus crenata Blume (Fagaceae) and Cryptomeria japonica D. Don (Cupressaceae). A flight mill was used to evaluate the effect of flight on the behavioral response to leaf volatiles. The bioassays were repeated 10× before and 10× after flight in the flight mill for each of the 54 individual beetles. Leaf volatile components were analyzed using gas chromatography–mass spectrometry. The bioassay results supported our hypothesis: P. quercivorus was attracted by the leaf volatiles of hosts and was deterred by the leaf volatiles of non-hosts. The behavioral response of P. quercivorus to the leaf volatiles was stronger after flight. Males had a stronger behavioral response than females to leaf volatiles. The leaf volatile chemical profile of the non-host C. japonica differed from the profile of the host plants. However, the chemical profile of the non-host F. crenata was similar to the profile of the hosts. Our findings provide insight into the functions of leaf chemical volatiles in the interaction of P. quercivorus with its hosts and non-hosts and may help improve the control of P. quercivorus and Japanese oak wilt.     

Fungus Symbionts Colonizing the Galleries of the Ambrosia Beetle <Emphasis Type="Italic">Platypus quercivorus</Emphasis>

Isolations were made to determine the fungal symbionts colonizing Platypus quercivorus beetle galleries of dead or dying Quercus laurifolia, Castanopsis cuspidata, Quercus serrata, Quercus crispula, and Quercus robur. For these studies, logs from oak wilt-killed trees were collected from Kyoto Prefecture, Japan. Fungi were isolated from the: (1) entrances of beetle galleries, (2) vertical galleries, (3) lateral galleries, and (4) the larval cradle of P. quercivorus in each host tree. Among the fungus colonies which appeared on YM agar plates, 1,219 were isolated as the representative isolates for fungus species inhabiting in the galleries based on their cultural characteristics. The validity of the visual classification of the fungus colonies was checked and if necessary properly corrected using microsatellite-primed PCR fingerprints. The nucleotide sequence of the D1/D2 region of the large subunit nuclear rRNA gene detected 38 fungus species (104 strains) of which three species, i.e., Candida sp. 3, Candida kashinagacola (both yeasts), and the filamentous fungus Raffaelea quercivora were isolated from all the tree species. The two yeasts were most prevalent in the interior of galleries, regardless of host tree species, suggesting their close association with the beetle. A culture-independent method, terminal restriction fragment length polymorphism (T-RFLP) analysis was also used to characterize the fungus flora of beetle galleries. T-RFLP patterns showed that yeast species belonging to the genus Ambrosiozyma frequently occurred on the gallery walls along with the two Candida species. Ours is the first report showing the specific fungi inhabiting the galleries of a platypodid ambrosia beetle.     

The effect of stem density on the probability of attack by the ambrosia beetle Platypus quercivorus varies with spatial scale

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An admixture of Quercus dentata in the coastal ecotype of Q. mongolica var. crispula in northern Hokkaido and genetic and environmental effects on their traits

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Species diversity estimation of ambrosia and bark beetles in temperate mixed forests in Japan based on host phylogeny and specificity

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Ambrosiozyma kamigamensis sp. nov. and A. neoplatypodis sp. nov., two new ascomycetous yeasts from ambrosia beetle galleries

Two new yeast species of the genus Ambrosiozyma are described on the basis of comparison of nucleotide sequences of large subunit of ribosomal DNA D1/D2 region. Ambrosiozyma kamigamensis and Ambrosiozyma neoplatypodis differ from Ambrosiozyma ambrosiae by 17 nucleotides (3.0%) and 16 nucleotides (2.8%), respectively, out of 565. The two species differ from each other by 13 nucleotides. Ambrosiozyma kamigamensis was isolated from galleries of the ambrosia beetle, Platypus quercivorus, in specimens of Quercus laurifolia and Castanopsis cuspidata located in the southern part of Kyoto, Japan. Ambrosiozyma neoplatypodis was isolated from similar material, but only in Q. laurifolia. Ambrosiozyma kamigamensis can be distinguished from the other Ambrosiozyma species by the inability to assimilate erythritol, whereas A. neoplatypodis can be distinguished by the ability to assimilate both L: -arabinose and nitrate. The type strains of A. kamigamensis and A. neoplatypodis are JCM 14990(T) (=CBS 10899(T)) and JCM 14992(T) (=CBS 10900(T)), respectively. This is the first report of new Ambrosiozyma species since the genus was proposed.     

Multiple evolutionary origins lead to diversity in the metabolic profiles of ambrosia fungi

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Phylogeographic structure and late Quaternary population history of the Japanese oak Quercus mongolica var. crispula and related species revealed by chloroplast DNA variation

Generally, oaks dominate the broadleaf deciduous forests in Japan. The genetic variation in 6 cpDNA regions (trnT-trnL, trnL-trnF, atpB-rbcL, and trnH-psbA speacers, trnL intron, and matK gene) with regard to the Japanese oak (Quercus mongolica var. crispula) and 3 related species in the section Prinus (Q. serrata, Q. dentata and Q. aliena) was investigated in 598 trees belonging to 44 populations distributed throughout the Japanese archipelago. Additional samples were collected from Korea, China, and Russia (Sakhalin). Thirteen haplotypes (I to XIII) were identified on the bases of 15 nucleotide substitutions and 3 indels. Haplotypes I and II were discovered in northeastern Japan, whereas haplotypes III to IX were distributed in southwestern Japan. The boundary distinguishing these 2 groups was located in central Japan coincident with the Itoigawa-Shzuoka tectonic line. Haplotype I was also found in Sakhalin, whereas haplotypes VI, VII, VIII, X, XI, XII, and XIII were found in Korea and China. Four oak species in the same location shared identical haplotypes, suggesting cpDNA introgression by occasional hybridization. Both the values of total haplotype diversity (HT) and haplotype diversity within populations (HS) in Q. mongolica var. crispula were higher in the southwestern populations than in the northeastern populations. A haplotype network indicated that haplotype VI is the ancestral haplotype. The presence of identical haplotypes in Korea, China, and Japan suggested that the haplotypes diversified on the Eurasian continent before the last glacial period. The difference in genetic structure between the northeastern and southwestern regions indicates a difference in the history of migration and recolonization in Japan during the last glacial period.     

Short-term development of ambrosia and bark beetle assemblages following a windstorm in French broadleaved temperate forests

Abstract: In most temperate deciduous forests, windstorm is the main source of dead wood. However, the effects of this natural disturbance on ambrosia and bark beetle communities are poorly known. In managed oak‐hornbeam forests storm‐damaged in France in 1999, we sampled ambrosia (and second bark beetles) by ethanol‐baited window‐flight traps in 2001. By comparing uncleared gaps, undisturbed closed‐canopy controls and seedling‐sapling stands, we investigated the short‐term effects of gap formation, gap size and surrounding landscape to provide a snapshot of scolytid response. Contrary to expectations, neither the abundance nor the richness of ambrosia beetle species was significantly higher in gaps than in undisturbed stands. Few responses in abundance at the species level and only a slight difference in assemblage composition were detected between gaps and closed‐canopy controls. Gaps were more dissimilar from seedling‐sapling stands, than from closed‐canopy controls. More scolytid individuals and species were caught in gaps than in seedling‐sapling stands. Mean local and cumulative richness peaked in mid‐size gaps. Only mid‐size gaps differed from closed‐canopy controls in terms of species composition. We identified generalist gap species (Xyleborus saxesenii, X. cryptographus), but also species significantly more abundant in mid‐size gaps (Platypus cylindrus, Xyloterus signatus). The faunistic peculiarity of mid‐size gaps seemed to be partly related to a bias in oak density among gap size classes. Few landscape effects were observed. Only the scolytids on the whole and X. dispar were slightly favoured by an increasing density in fellings at the 78 ha scale. We did not find any correlation between scolytid abundance and the surrounding closed‐forest percentage area. We confirmed that temperate, deciduous, managed stands did not come under threat by ambrosia and bark beetle pests after the 1999 windstorm. Nonetheless, our data stressed the current expansion in Western Europe of two invasive species, X. peregrinus and especially X. germanus, now the predominant scolytid in the three oak forests studied.     

Identification of the ambrosia fungus of <Emphasis Type="Italic">Xyleborus monographus</Emphasis> and <Emphasis Type="Italic">X. dryographus</Emphasis> (Coleoptera: Curculionidae,Scolytinae)

The scolytid ambrosia beetles Xyleborus monographus and X. dryographus were investigated to identify their nutritional ambrosia fungi. The examination of the oral mycetangia of the beetles, the specialized organs for fungal transport, revealed the dominant occurrence of Raffaelea montetyi, a fungus that was also predominant in the beetle tunnels in the immediate vicinity of the feeding larvae. R. montetyi was previously known only as the ambrosia fungus of the platypodid ambrosia beetle, Platypus cylindrus. These beetle species inhabit the same habitat, mainly trunks of oaks in the Western Palaeartic. The possibility of an exchange of the symbiotic fungus between the ambrosia beetles within their common breeding place is discussed. Consequently, the previous hypothesis of a species-specific association of a single ambrosia fungus with a single beetle species is questioned. A phylogenetic analysis based on DNA sequences classified R. montetyi within the Ophiostomatales of the ascomycetes. The investigation of conidiogenesis of R. montetyi by SEM supported this taxonomic placement and showed the development of the conidia by annellidic percurrent proliferation, identical to the conidiogenesis reported for many anamorph states of the Ophiostomatales.     

External microstructure of the ambrosia beetle Platypus koryoensis (Coleoptera: Curculionidae: Platypodinae)

Platypus koryoensis is a minute ambrosia beetle found in forests. It can cause significant economic damage to oak trees. Recently in Korea, it has been reported as a major pest of oak trees, because it causes sooty mold of oak by introducing the pathogenic fungus Raffaelea sp. In this paper, we demonstrate the fine structural aspects of the external body of the ambrosia beetle using field emission scanning electron microscopy, as a part of basic research into this pest so that strategies for its control might be developed. This beetle has a sensory system well developed to respond to both visual and chemical stimuli. Both sexes have a pair of faceted compound eyes and a pair of knobbed antennae, but simple eyes are absent. The mouthparts on its distinct snouts are effective devices for penetration and for boring holes. The mouthparts consist of the labrum, a pair of mandibles, a pair of maxillae and the labium. Both the maxillary and the labial palpi have the function of directing the food to the mouth and holding it while the mandibles chew the food. The distal ends of these palpi are flattened and have shovel‐like setae. The thorax has a particularly hard exoskeleton and hard elytra, including powerful muscles that operate both the wings and the legs. The legs are multi‐segmented and have a strong femur and tibia, including one pair of claws on the end of each tarsal segment. Characteristically, both male and female beetles have mycangial cavities for storing spores and other microorganisms, but only females have three pairs of large depressions on their dorsal thorax.     

Microstructure of the prothoracic mycangia in the ambrosia beetle Platypus koryoensis (Coleoptera: Curculionidae: Platypodinae)

Platypus koryoensis (Murayama) (Coleoptera: Curculionidae: Platypodinae) is a minute ambrosia beetle that can cause significant damage to live oak trees (Quercus mongolica) along with its symbiotic pathogenic fungus (Raffaelea sp.). Although both sexes of the beetle have a number of cuticular depressions for carrying microorganisms, only the females have mycangia that are typical in size on the pronotum. The mycangial cavities are filled with a mass of amorphous substances including yeast-like spores. In addition, mycangial cavities are equipped with cuticular pores which release substances to support symbiotic microorganisms during storage. Small depressions of both sexes can be subdivided into two clusters according to their relative sizes and locations. Although they have a typical cuticular peg (basiconic sensillum), they also share a common morphological characteristic with the typical mycangium of female: a round outlet and an invaginated reservoir containing a mass of secretory substances. Accordingly, our fine structural data suggest that both the female mycangia and the clusters of small cuticular depressions in both sexes also have a mycangial function in common regardless of their relative size and location.     

Attack pattern of Platypus koryoensis (Coleoptera: Curculionidae: Platypodinae) in relation to crown dieback of Mongolian oak in Korea

The ambrosia beetle, Platypus koryoensis (Murayama), vectors the Korean oak wilt (KOW) pathogen, Raffaelea quercus-mongolicae K.H. Kim, Y.J. Choi, & H.D. Shin, in Korea, which is highly lethal to Mongolian oak, Quercus mongolica Fisch., and is considered a major threat to forest ecosystem health. We characterized the attack pattern of P. koryoensis along the lower trunk of 240 Mongolian oaks in relation to tree decline symptoms on Mt. Uam in Gyeonggi-Do Province, Korea during June-July 2009. For each tree, we recorded diameter at breast height (dbh) (DBH) and P. koryoensis entrance hole density at two heights along the lower trunk (near groundline and at 1.5 m above groundline) and on opposite sides (downslope side and upslope side). Trees were assigned to one of three dieback classes: 1) apparently healthy, no or practically no wilted foliage, and no obvious platypodine frass near the base of the tree; 2) no or only partial wilting with obvious frass near the base of the tree; and 3) apparently recently killed by KOW with all foliage wilted and mostly retained with obvious frass near the base of the tree. As dieback class increased from 1 to 3, P. koryoensis entrance hole density increased at all four trunk locations. Attack density was highest on the downslope side of the trunk near groundline, and principal component analysis indicated that this trunk location was the best indicator of tree dieback. In addition, DBH tended to increase with dieback class suggesting that larger trees were infested first.     

The influence of forest stand and site characteristics on the composition of exotic dominated ambrosia beetle communities (Coleoptera: Curculionidae: Scolytinae)

Economic and biological consequences are associated with exotic ambrosia beetles and their fungal associates. Despite this, knowledge of ambrosia beetles and their ecological interactions remain poorly understood, especially in the oak-hickory forest region. We examined how forest stand and site characteristics influenced ambrosia beetle habitat use as evaluated by species richness and abundance of ambrosia beetles, both the native component and individual exotic species. We documented the species composition of the ambrosia beetle community, flight activity, and habitat use over a 2-yr period by placing flight traps in regenerating clearcuts and older oak-hickory forest stands differing in topographic aspect. The ambrosia beetle community consisted of 20 species with exotic ambrosia beetle species dominating the community. Similar percentages of exotic ambrosia beetles occurred among the four forest habitats despite differences in stand age and aspect. Stand characteristics, such as stand age and forest structure, influenced ambrosia beetle richness and the abundances of a few exotic ambrosia beetle species and the native ambrosia beetle component. Topographic aspect had little influence on ambrosia beetle abundance or species richness. Older forests typically have more host material than younger forests and our results may be related to the amount of dead wood present. Different forms of forest management may not alter the percent contribution of exotic ambrosia beetles to the ambrosia beetle community.