Effects of leaf conditions and flight activity on the behaviour of Platypus quercivorus (Murayama) (Coleoptera: Platypodidae)

The important role of semiochemicals in the interactions between plants and insects has been extensively investigated. The volatiles produced by oak trees are thought to attract the ambrosia beetle Platypus quercivorus (Murayama), which causes wilt disease in Quercus trees, resulting in widespread damage. In the present study, we hypothesized that (a) P. quercivorus is attracted to the leaf volatiles emitted by host trees and (b) the response of P. quercivorus to leaf volatiles is affected by flight. An experiment was performed to survey the preferences of both sexes of P. quercivorus for the leaf volatiles of Quercus crispula Blume at various stages of leaf deterioration, represented by different number of days after cutting. Additionally, the effect of flight on both sexes was evaluated by testing the beetle flight on a flight mill. The results showed that P. quercivorus was attracted to the volatiles emitted from fresh leaves and was not attracted to those emitted from dry leaves. This suggests that leaf volatiles from healthy hosts are primary attractants for P. quercivorus. Further, males exhibited increased olfactory responses to leaf volatiles as their flight duration increased, whereas the opposite pattern was observed in females. These results suggest that the different ecological roles of the sexes contribute to differences in olfactory responses.     

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

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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     

Additive and synergistic integration of multimodal cues of both hosts and non-hosts during host selection by woodboring insects

Host selection by herbivorous insects is primarily thought to depend on attraction to olfactory cues emitted from the host species. However, the discrimination of these hosts from non-host species may also arise from the adaptive detection and avoidance of non-host cues, including visual characteristics. Many generalist, conifer-colonising beetles, for example, use characteristic volatiles to identify and discriminate against non-host angiosperm trees such as aspens and birches while flying. These trees also differ in bark reflectance characteristics, which could also mediate host/non-host discrimination by interacting with semiochemicals. We tested this hypothesis by evaluating the responses of eight species of polyphagous woodboring beetles to traps which simulated the visual appearance of coniferous hosts (black) and angiosperm non-hosts (white), and which were baited with host or non-host volatiles. As predicted, three species of large woodboring beetle and a woodboring wasp all avoided white, non-host-simulating traps that were baited with attractive kairomones, and preferred black, host-simulating traps. Conversely, three ambrosia beetle species demonstrated weaker visual preferences, possibly because they preferentially colonise fallen hosts that would transmit less accurate visual information. However, the ambrosia beetle Trypodendron lineatum did show a greater preference for host-coloured traps when these released host-associated kairomones in addition to their pheromone, and also avoided white non-host traps, but only when these released non-host volatiles. To our knowledge, this is the first evidence that multiple non-host cues could synergistically mediate the adaptive discrimination of hosts and non-hosts. Our results suggest that successful host location by generalists arises from the complex integration of cues in multiple sensory modes, and that foraging herbivores evaluate both hosts and non-hosts during their search.     

Phototactic behavior of the ambrosia beetle <Emphasis Type="Italic">Platypus quercivorus</Emphasis> (Murayama) (Coleoptera: Platypodidae) before and after flight

The ambrosia beetle Platypus quercivorus (Coleoptera: Platypodidae) is a vector of the pathogenic fungi, Raffaelea quercivora (Ophiostomatales: Ophiostomataceae), which causes Japanese oak wilt disease. Previous studies have shown that the beetle displays positive phototactic behavior; however, the adaptive significance of this behavior remains unclear. We postulated that positive phototaxis is necessary to allow the beetle to fly skyward immediately after emergence from a tree, and that this taxis changes following a certain period of flight. The present study aimed to clarify the changes in phototactic behavior of P. quercivorus before and after flight by using individual beetles emerging from the trunk of a Quercus crispula (Fagales: Fagaceae) tree that was attacked in 2014. The response of 60 beetles to light was tested ten times each, before and after flight in a flight mill. A generalized linear mixed model was constructed to predict the probability of positive phototaxis of P. quercivorus before and after flight. A best-fit model showed that the probability of positive phototaxis was lower after flight than before. The results suggest that positive phototaxis of P. quercivorus is decreased after flight.     

Shoot structure and dynamics of saplings and canopies of three deciduous broad-leaved trees of a coppice forest in central Japan

Three deciduous broad-leaved trees, Quercus serrata, Castanea crenata and Carpinus laxiflora, were the main constituents of a coppice forest in central Japan. The shoot elongation and leaf emergence modes of both saplings and the canopy of the three species were investigated. The shoot elongation modes of Q. serrata and C. crenata were the same in saplings and the upper layer of the canopy. The second shoots of these two species were formed after the first shoots were elongated. C. laxiflora was different between saplings and the upper layer of the canopy. In saplings, only the first shoots took a long time to elongate. In the upper canopy layer, higher order shoots were formed in the same way as in the other two species. In the lower layer of the canopy, all three species showed the same shoot elongation mode, in which only the first shoot and its duration of elongation was short. Leaf longevity, individual leaf area, leaf mass per unit leaf area and the stem mass per unit stem length of C. laxiflora were significantly shorter or significantly smaller than those of Q. serrata and C. crenata. The length of the stem per unit leaf area of C. laxiflora was three times that of Q. serrata and five times that of C. crenata. The elongation growth of C. laxiflora was highly efficient as it occurred with a small leaf area. The shoot dynamics and the shoot structure of C. laxiflora are more suitable for elongation growth than in Q. serrata and C. crenata. Furthermore, the shoot structures of the three species were compared and ecological characteristics of the three species are discussed. Received: 29 September 1998 / Accepted: 17 September 1999     

Stand-level distribution and movement of Platypus quercivorus adults and patterns of incidence of new infestation

Abstract 1 Flying populations of an ambrosia beetle, Platypus quercivorus (Murayama), a vector of an ambrosia fungus Raffaelea quercivora, which causes deciduous oak diebacks in Japan, were determined by sticky screen traps. 2 Platypus quercivorus beetles tended to move upwards along slopes. The highest concentrations of flying beetles usually occured at the upper forest margins. 3 During the period when the number of flying beetles was increasing, the incidence of newly infested trees spread from the epicentre into the forest. During the period when the number of flying beetles was decreasing, the epicentre shrank into the upper forest edge. Newly infested trees did not occur in this period because most trees had already been infested. 4 Near the upper forest edge, where many beetles were highly concentrated throughout the season, the number of new entry holes decreased greatly after the initial attack early in the season, although many adults were present throughout the period.     

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.     

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.     

Plastic leaf morphology in three species of Quercus: The more exposed leaves are smaller,more lobated and denser

Phenotypic plasticity and developmental instability in leaf traits are common in oak species but the role of environmental factors is not well understood. To decipher possible correlations between different leaf traits and effects of the position of leaves within the tree canopy, we quantified the plasticity of three leaf traits of 30 trees of Quercus alba L., Quercus palustris Muench and Quercus velutina Lam. We hypothesized that trees could modify the shape of their leaves for better adaptation to the variable microclimate within the canopy. Our results demonstrated that the south and north outer leaves were significantly smaller, more lobed and denser than those situated in the inner canopy. The order of leaves on the branch accounted for the plasticity of leaf traits in Q. alba only. Plasticity of lobing in Q. alba and Q. velutina depended on the height of the trees. We detected fluctuating asymmetry (FA) in all three species, but the source of variation depended on branch position in Q. velutina only. FA was more pronounced in north-facing leaves. Plasticity of the leaf traits ranged from small to medium. Plasticity of leaf area and leaf mass per area (LMA) depended on the branch position. However, the plasticity of lobation was not affected by the location of a branch within the tree canopy. Quercus alba and Q. palustris had similar plastic responses but the plasticity of Q. velutina was significantly smaller. We concluded that individual plants detect and cope with environmental stress through vegetative organ modification.     

Potential impact of global warming on deciduous oak dieback caused by ambrosia fungus Raffaelea sp. carried by ambrosia beetle Platypus quercivorus (Coleoptera: Platypodidae) in Japan

Deciduous oak dieback in Japan has been known since the 1930s, but in the last ten years epidemics have intensified and spread to the island's western coastal areas. The symbiotic ambrosia fungus Raffaelea sp. is the causal agent of oak dieback, and is vectored by Platypus quercivorus (Murayama). This is the first example of an ambrosia beetle fungus that kills vigorous trees. Mortality of Quercus crispula was approximately 40% but much lower for associated species of Fagaceae, even though each species had a similar number of beetle attacks. It is likely that other oaks resistant to the fungus evolved under a stable relationship between the tree, fungus and beetle during a long evolutionary process. Quercus crispula was probably not part of this coevolution. This hypothesis was supported by the fact that P. quercivorus showed the least preference for Q. crispula yet exhibited highest reproductive success in this species. Therefore, P. quercivorus could spread more rapidly in stands with a high composition of Q. crispula. The present oak dieback epidemic in Japan probably resulted from the warmer climate that occurred from the late 1980s which made possible the fateful encounter of P. quercivorus with Q. cripsula by allowing the beetle to extend its distribution to more northerly latitudes and higher altitudes. Future global warming will possibly accelerate the overlapping of the distributions of P. quercivorus and Q. crispula with the result that oak dieback in Q. crispula will become more prevalent in Japan.     

Spread and potential host range of the invasive oak lace bug [Corythucha arcuata (Say, 1832) – Heteroptera: Tingidae] in Eurasia

1. The North American oak lace bug feeds on leaves of ‘white oaks” in its native range. In Europe, it was first discovered in northern Italy in 2000. In recent years, it has subsequently spread rapidly and population outbreaks have been observed in several European countries. In the present study, we summarize the steps of its expansion.
2. To predict its potential host range, we checked 48 oak species in 20 sentinel gardens in seven countries between 2013 and 2018.
3. In total, 27 oak species were recorded as suitable hosts; 13 of them are globally new ones, 23 out of the 29 in section Quercus (～ white oaks, an intrageneric taxonomic unit within genus Quercus), including Asian oaks, native to Japan, Korea and China, and four out of five in section Cerris (another intrageneric unit of the same genus), were accepted as hosts. None of the species in section Lobatae (red oaks) or in the Ilex group was accepted.
4. Host records were also collected in forest stands of 10 countries. We found 11 oak species that were infested. Outbreak populations were most commonly found on Quercus robur, Quercus frainetto, Quercus petraea and Quercus cerris, comprising widespread and outstandingly important oaks species in Europe.
5. Based on our findings, we conclude that suitable hosts for oak lace bug are present in most of Europe and Asia. This means that a lack of hosts will likely not restrict further range expansion.

     

Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

Didymella fagi sp. nov. and its anamorphAscochyta fagi, causing the yellow leaf spot disease ofFagus crenata andQuercus mongolica var.grosseserrata in Japan

AnAscochyta fungus was obtained during a survey of leaf spot diseases ofFagus crenata in the mountains of Aomori prefecture in 1995. The pathogenicity of the fungus toF. crenata was confirmed by inoculation. Its teleomorph was first found on artificially infected leaves after the leaves were placed in an incubator at 5°C with a 12-h photoperiod (approximately 500lx, daylight strip lamps) for 4 mo. The fungus was found to be the causal agent of yellow leaf spots ofF. crenata andQuercus mongolica var.grosseserrata in the field. Comparison with similar fungi so far described from Fagaceae indicated that the anamorph isAscochyta fagi and the teleomorph is an undescribed species ofDidymella. The nameDidymella fagi is introduced for the teleomorph. Correlation between the two morphs has been proved by cultural, morphological and pathological studies.     

Pathogenicity of microorganisms isolated from the oak platypodid, <Emphasis Type="Italic">Platypus quercivorus</Emphasis> (Murayama) (Coleoptera: Platypodidae)

The oak platypodid, Platypus quercivorus (Murayama), is the most destructive pest of Quercus and related fagaceous species in Japan. To find an effective biocontrol agent, we determined the effectiveness of entomopathogenic microorganisms against P. quercivorus. P. quercivorus-infested Oak (Quercus serrata, Q. crispula and Q. acutissima) logs were obtained from various regions in Japan, and 28 isolates of microorganisms were isolated from P. quercivorus larvae from those logs. Using molecular methods, the microbes were then classified into 18 species. After a bioassay, we found that three fungi, Lecanicillium sp., Isaria tenuipes, Beauveria bassiana, and the bacterium Serratia marcescens had the greatest potential as biological control agents against P. quercivorus. Another bioassay was then performed to determine the effect of spore or cell densities on the pathogenicity of the four microorganisms, and optimum spore or cell densities for the microbes were determined to be 10⁶ and 10⁷/ml. Based on the results and other criteria, we tentatively selected B. bassiana as the best candidate for future use against P. quercivorus larvae.     

Tree species identity and forest composition affect the number of oak processionary moth captured in pheromone traps and the intensity of larval defoliation

1. Pure forests are often seen as being more prone to damage by specialist pest insects than mixed forests, and particularly mixed forests associating host and nonhost species. We addressed the effect of tree diversity on oak colonization and defoliation by a major specialist pest, the oak processionary moth (OPM)
2. We quantified the number of male OPM moths captured and larval defoliation in pure stands of two oak host species (Quercus robur and Quercus petraea) and in mixed stands associating the two oak species or each oak species with another nonhost broadleaved species. We conducted two complementary studies to test the effect of host species and stand composition: (i) we used pheromone trapping to compare the number of males OPM captured throughout the distribution of oak hosts in France and (ii) we noted the presence of OPM nests and estimated defoliation in mature forests of north‐eastern France.
3. Oak species and stand composition significantly influenced the number of male OPM captured and defoliation by OPM larvae. Quercus petraea was consistently more attractive to and more defoliated by OPM than Q. robur. Both oak trees were attacked more in pure stands than in mixed stands, in particular mixed stands associating oaks with another (nonhost) broadleaved species.
4. The results of the present study support the view that mixed forests are more resistant to specialist pest insects than pure stands, and also indicate that this trend depends on forest composition. Our study provides new insights into OPM ecology and has potential implications for forest management, including the management of urban forests where OPM causes serious human health issues.

     

Growth and maintenance respiration in leaves of northern red oak seedlings and mature trees after 3 years of ozone exposure

A two-component model of growth and maintenance respiration is used to study the response of northern red oak (Quercus rubra L.) seedlings and 32-year-old trees to sub-ambient (10 μmol h; cumulative dose based on 7 h daily mean), ambient (43 μmol h), and twice-ambient (85 μmolh) ozone. The relative growth rates (RGR) of leaves sampled from seedlings and trees were similar across treatments, as were specific leaf respiration rates (SRR). Growth coefficients estimated from the SRR versus RGR relationship averaged 25-3 mol CO2 kg^?1 leaf dry mass produced for seedlings and 21-5 mol kg^?1 for trees. Maintenance coefficients ranged from 0-89 to 1-07 mol CO₂ kg^?1 leaf dry mass d^?1 for seedlings and from 0-64 to 0-84 mol kg-1 d^?1 for trees. Neither coefficient was affected by ozone. Leaves sampled throughout the growing season also showed little response of respiration to ozone. This occurred despite a 30% reduction in net photosynthesis for trees grown at twice-ambient ozone. These results suggest that growth and maintenance respiration in young northern red oak leaves are not affected by ozone and that in older leaves injury can occur without a parallel increase in so-called ‘maintenance’ respiration.     

Foliar nutrients explain goldspotted oak borer,Agrilus auroguttatus,adult feeding preference among four California oak species

Adults of the invasive goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), consumed foliar weight in no‐choice feeding tests of, in descending order, California black oak Quercus kelloggii Newb., Engelmann oak, Quercus engelmannii Greene, coast live oak, Quercus agrifolia Née, and canyon live oak, Quercus chrysolepis Liebm. (Fagaceae). Furthermore, significantly more foliar area was consumed of Q. kelloggii than of Q. chrysolepis. In dual‐choice feeding tests with isolated leaf disks, A. auroguttatus consumed significantly more foliar weight and area of Q. kelloggii relative to the other three oak species, and more foliar weight of Q. agrifolia than of Q. chrysolepis. In dual‐choice feeding tests with leaves on small branches, A. auroguttatus consumed more foliar weight of Q. kelloggii than of Q. engelmannii and Q. agrifolia. Thus, multiple experiments suggested that adults of A. auroguttatus preferred the foliage of Q. kelloggii over that of the other three oak species, and among the other three species they did not appear to have a strong feeding preference. Factor analysis reduced the quantities of 13 foliar nutrients into two new variables (factor 1 and factor 2). Factor 1 was weighted heavily on the quantities of nitrogen, sulfur, phosphorus, potassium, zinc, and copper, whereas factor 2 was weighted heavily on the quantities of zinc, iron, and aluminum. Factor 1 varied by oak species, with Q. kelloggii having a higher factor 1 nutrient content than the other three species. Factor 2 response was higher in Q. kelloggii, Q. agrifolia, and Q. engelmannii than in Q. chrysolepis. The collective effects of four macronutrients (nitrogen, sulfur, phosphorus, and potassium) and two micronutrients (zinc and copper) suggest that these might be the nutrients directing preferential feeding of A. auroguttatus adults on the foliage of Q. kelloggii. Leaf toughness might also play an important role in feeding preference. Female A. auroguttatus did not show an ovipositional preference among the four oak species.