Invasive bark beetle‐associated microbes degrade a host defensive monoterpene

Conifers respond to herbivore attack with defensive chemicals, which are toxic to both insects and their associated microorganisms. Microorganisms associated with insects have been widely reported to metabolize toxic chemicals, which may help both microorganisms and host insects overcome host conifer defense. Dendroctonus valens LeConte, an introduced exotic pest from North America to China, has killed millions of healthy pines. Alpha‐pinene is the most abundant defensive monoterpene in Chinese Pinus tabuliformis. Although microorganisms associated with D. valens have already been investigated, little is known about their bioactivities when encountering host defensive monoterpenes. In this study, we evaluated the influences of different concentrations of α‐pinene to D. valens and the three most frequently isolated yeasts and bacteria of D. valens, and further assayed microorganisms’ capabilities to degrade α‐pinene. Results showed that the gallery lengths and body weight changes of bark beetles were significantly affected by 6 mg/mL and 12 mg/mL of α‐pinene applied in media compared to controls. The tolerance of experimental microorganisms to α‐pinene varied depending on the microbial species. Two out of three yeast strains and all three bacterial strains degraded 20%–50% of α‐pinene compared to controls in 24 h in vitro. The microorganisms capable of α‐pinene degradation in vitro and their tolerance to high levels of α‐pinene suggested that D. valens‐associated microorganisms may help both microorganisms and the bark beetle overcome host α‐pinene defense.     

Attraction to monoterpenes and beetle‐produced compounds by syntopic Ips and Dendroctonus bark beetles and their predators

• 1 Bark beetles are significant mortality agents of conifers. Four beetle species, the pine engraver Ips pini, the six‐spined pine engraver Ips calligraphus sub. ponderosae, the southern pine beetle Dendroctonus frontalis, and the western pine beetle Dendroctonus brevicomis, cohabitate pines in Arizona.
• 2 A pheromone trapping study in ponderosa forests of Arizona determined the attraction of beetles to conspecific and heterospecific pheromone components in the presence and absence of host volatiles, and tested whether predators differ in their attraction to combinations of pheromone components and tree monoterpenes.
• 3 All four bark beetle species differed in their responses to heterospecific lures and monoterpenes. Ips calligraphus was the only species that increased in trap catches when heterospecific lures were added. Heterospecific lures did not inhibit the attraction of either Dendroctonus or Ips species. The replacement of myrcene with α‐pinene increased the attraction of Dendroctonus, whereas the addition of α‐pinene had mixed results for Ips. The prominent predators Temnochila chlorodia and Enoclerus lecontei were more attracted to the I. pini lure than the D. brevicomis lure, and the combination of the two lures with α‐pinene was most attractive to both predator species.
• 4 Cross attraction and limited inhibition of bark beetles to heterospecific pheromones suggest that some of these species might use heterospecific compounds to increase successful location and colonization of trees. Predator responses to treatments suggest that tree volatiles are used to locate potential prey and predators are more responsive to Ips than to Dendroctonus pheromone components in Arizona.

     

Synergistic effects of α-pinene and exo-brevicomin on pine bark beetles and associated insects in Arizona

The southern pine beetle (Dendroctonus frontalis) and western pine beetle (Dendroctonus brevicomis) cause significant mortality to pines in the southern and western United States. The effectiveness of commercial lures at capturing these bark beetles in Arizona has not been tested and may vary from other regions of their distribution. We conducted experiments using baited Lindgren funnel traps to investigate (i) if D. frontalis is more attracted to the standard commercial lure for D. brevicomis (frontalin + exo‐brevicomin + myrcene) than the D. frontalis lure (frontalin + terpene blend), (ii) whether replacement of myrcene with α‐pinene changes trap catches of Dendroctonus and associated insects, and (iii) whether the attraction to these lures varies across the geographical range of ponderosa pine forests throughout Arizona. In 2005, we tested various combinations of frontalin, exo‐brevicomin, myrcene and α‐pinene to D. frontalis, D. brevicomis and associated species. Dendroctonus frontalis, D. brevicomis and the predator Temnochila chlorodia were most attracted to lures with exo‐brevicomin. The replacement of the myrcene component with α‐pinene in the D. brevicomis lure resulted in the capture of twice as many bark beetles and Elacatis beetles. However, T. chlorodia did not differentiate between monoterpenes. In 2006, traps were set up in 11 locations around Arizona to test the relative attraction of lure combinations. In 9 out 11 locations, the D. brevicomis lure with α‐pinene was more attractive than the lure with myrcene or a terpene blend. These results suggest that the D. brevicomis lure with α‐pinene rather than myrcene is more effective lure to capture D. brevicomis and D. frontalis in Arizona. However, geographical variation in attractiveness to lures is evident even within this region of the beetles’ distributions. Differential attraction of Dendroctonus and their predators to these lures suggests potential use in field trapping and control programmes.     

2‐(Undecyloxy)‐ethanol is a major component of the male‐produced aggregation pheromone of Monochamus sutor

The small white‐marmorated longicorn beetle, Monochamus sutor (L.) (Coleoptera: Cerambycidae), is widely distributed throughout Europe and Asia. It is a potential vector of the pine wood nematode, Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle, the causal agent of the devastating pine wilt disease. Volatiles were collected from both male and female beetles after maturation feeding. In analyses of these collections using gas chromatography (GC) coupled to mass spectrometry, a single male‐specific compound was detected and identified as 2‐(undecyloxy)‐ethanol. In analyses by GC coupled to electroantennography the only consistent responses from both female and male antennae were to this compound. Trapping tests were carried out in Spain, Sweden, and China. 2‐(Undecyloxy)‐ethanol was attractive to both male and female M. sutor beetles. A blend of the bark beetle pheromones ipsenol, ipsdienol, and 2‐methyl‐3‐buten‐2‐ol was also attractive to both sexes in Spain and Sweden, and further increased the attractiveness of the 2‐(undecyloxy)‐ethanol. The host plant volatiles α‐pinene, 3‐carene, and ethanol were weakly attractive, if at all, in all three countries and did not significantly increase the attractiveness of the blend of 2‐(undecyloxy)‐ethanol and bark beetle pheromones. 2‐(Undecyloxy)‐ethanol is thus proposed to be the major, if not only, component of the male‐produced aggregation pheromone of M. sutor, and its role is discussed. This compound has been reported as a pheromone of several other Monochamus species and is another example of the parsimony that seems to exist among the pheromones of many of the Cerambycidae. Traps baited with 2‐(undecyloxy)‐ethanol and bark beetle pheromones should be useful for monitoring and control of pine wilt disease, should M. sutor be proven to be a vector of the nematode.     

The red turpentine beetle, <Emphasis Type="Italic">Dendroctonus valens</Emphasis> LeConte (Scolytidae): an exotic invasive pest of pine in China

An exotic invasive pest of pines, the red turpentine beetle, Dendroctonus valens LeConte (Scolytidae) (RTB), was first detected in Shanxi Province, northern China, in 1998 and started causing widespread tree mortality there in 1999. This outbreak continues and has spread to three adjacent provinces, causing unprecedented tree mortality. Although it is considered a minor pest of pines in North America, RTB has proven to be an aggressive and destructive pest of Pinus tabuliformis, China’s most widely planted pine species. The bionomics and occurrence, distribution, response to host volatiles, and host preference of this pine beetle in China are compared with what is known of the beetle in its native range in North America. Factors likely contributing to D. valens success in China and control of the beetle outbreak are discussed. (−)-β-pinene was shown to be the most attractive host volatile for D. valens from the Sierra Nevada of California, whereas 3-(+)-carene is the most attractive host volatile for beetles in China. Monocultures of Pinus tabuliformis, several consecutive years of drought conditions and warm winters have apparently factored D. valens invasion and establishment in China.     

Attraction of red turpentine beetle and other Scolytinae to ethanol, 3‐carene or ethanol + 3‐carene in an Oregon pine forest

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Effect of varying monoterpene concentrations on the response of Ips pini (Coleoptera: Scolytidae) to its aggregation pheromone: implications for pest management and ecology of bark beetles

Abstract 1 Host plant terpenes can influence attraction of conifer bark beetles to their aggregation pheromones: both synergistic and inhibitory compounds have been reported. However, we know little about how varying concentrations of individual monoterpenes affect responses. 2 We tested a gradient of ratios of α‐pinene, the predominant monoterpene in host pines in the Great Lakes region of North America, to Ips pini's pheromone, racemic ipsdienol plus lanierone. 3 Ips pini demonstrated a parabolic response, in which low concentrations of α‐pinene had no effect on attraction to its pheromone, intermediate concentrations were synergistic and high concentrations were inhibitory. These results suggest optimal release rates for population monitoring and suppression programmes. 4 Inhibition of bark beetle attraction to pheromones may be an important component of conifer defences. At terpene to pheromone ratios emulating emissions from trees actively responding to a first attack, arrival of flying beetles was low. This may constitute an additional defensive role of terpenes, which are also toxic to bark beetles at high concentrations. 5 Reduced attraction to a low ratio of α‐pinene to pheromone, as occurs when colonization densities become high and the tree's resin is largely depleted, might reflect a mechanism for preventing excessive crowding. 6 Thanasimus dubius, the predominant predator of I. pini, was also attracted to ipsdienol plus lanierone, but its response differed from that of its prey. Attraction increased across all concentrations of α‐pinene. This indicates that separate lures are needed to sample both predators and bark beetles effectively. It also provides an opportunity for maximizing pest removal while reducing adverse effects on beneficial species. This disparity further illustrates the complexity confronting natural enemies that track chemical signals to locate herbivores.     

Cover Caption

Dendroctonus valens, an introduced exotic pest from North America to China, has killed millions of healthy Chinese pines. Xu et al. (2016) showed two yeasts and three bacteria associated with the invasive pest degraded 20%‐50% of 〈‐pinene, the most abundant defensive monoterpene of Chinese pines. The microorganisms capable of 〈‐pinene degradation in vitro and their tolerance to high levels of 〈‐pinene suggested that D. valens‐associated microorganisms may help both microorganisms and the bark beetle overcome host 〈‐pinene defense (see pages 183‐190). Photo by Run‐Zhi Zhang.     

Selective manipulation of predators using pheromones: responses to frontalin and ipsdienol pheromone components of bark beetles in the Great Lakes region

Abstract 1 One proposed approach to improving biological control of bark beetles (Coleoptera: Scolytidae; alt. Curculionidae: Scolytinae) is to manipulate predator movement using semiochemicals. However, selective manipulation is impeded by attraction of both predators and pests to bark beetle pheromones. 2 The primary bark beetle affecting pine plantations in Wisconsin, U.S.A., is the pine engraver, Ips pini (Say). Other herbivores include Ips grandicollis (Eichhoff) and Dryophthorus americanus Bedel (Curculionidae). The predominant predators are the beetles Thanasimus dubius (Cleridae) and Platysoma cylindrica (Histeridae). 3 We conducted field assays using two enantiomeric ratios of ipsdienol, and frontalin plus α‐pinene. Ipsdienol is the principal pheromone component of I. pini, and frontalin is produced by a number of Dendroctonus species. α‐Pinene is a host monoterpene commonly incorporated into commercial frontalin lures. 4 Thanasimus dubius was attracted to frontalin plus α‐pinene, and also to racemic ipsdienol. By contrast, I. pini was attracted to racemic ipsdienol, but showed no attraction to frontalin plus α‐pinene. Platysoma cylindrica was attracted to 97%‐(–)‐ipsdienol and, to a lesser extent, racemic ipsdienol, but not to frontalin plus α‐pinene. Ips grandicollis was attracted to frontalin plus α‐pinene but not to ipsdienol. Dryophthorus americanus was attracted to both ipsdienol and frontalin plus α‐pinene. 5 This ability to selectively attract the predator T. dubius without attracting the principal bark beetle in the system, I. pini, provides new opportunities for research into augmentative biological control and basic population dynamics. Moreover, the attraction of T. dubius, but not P. cylindrica, to frontalin plus α‐pinene creates opportunities for selective manipulation of just one predator. 6 Patterns of attraction by predators and bark beetles to these compounds appear to reflect various degrees of geographical and host tree overlap with several pheromone‐producing species.     

Cross-Attraction between an Exotic and a Native Pine Bark Beetle: A Novel Invasion Mechanism?

Background

Aside from the ecological impacts, invasive species fascinate ecologists because of the unique opportunities that invasives offer in the study of community ecology. Some hypotheses have been proposed to illustrate the mechanisms that allow exotics to become invasive. However, positive interactions between exotic and native insects are rarely utilized to explain invasiveness of pests.

Methodology/Principal Findings

Here, we present information on a recently formed association between a native and an exotic bark beetle on their shared host, Pinus tabuliformis, in China. In field examinations, we found that 35–40% of P. tabuliformis attacked by an exotic bark beetle, Dendroctonus valens, were also attacked by a native pine bark beetle, Hylastes parallelus. In the laboratory, we found that the antennal and walking responses of H. parallelus to host- and beetle-produced compounds were similar to those of the exotic D. valens in China. In addition, D. valens was attracted to volatiles produced by the native H. parallelus.

Conclusions/Significance

We report, for the first time, facilitation between an exotic and a native bark beetle seems to involve overlap in the use of host attractants and pheromones, which is cross-attraction. The concept of this interspecific facilitation could be explored as a novel invasive mechanism which helps explain invasiveness of not only exotic bark beetles but also other introduced pests in principle. The results reported here also have particularly important implications for risk assessments and management strategies for invasive species.     

Field evaluation on the synergistic attractiveness of 2‐(1‐undecyloxy)‐1‐ethanol and ipsenol to Monochamus saltuarius

To develop an optimal attractant for Monochamus saltuarius (Gebler) (Coleoptera: Cerambycidae), the synergistic effects of a few potential attractants (ethanol and α‐pinene as host‐plant volatiles, and ipsenol and ipsdienol as bark beetle pheromones) were tested in a pine forest combined with 2‐(1‐undecyloxy)‐1‐ethanol (monochamol), the aggregation pheromone of Monochamus species, for two consecutive years, 2014 and 2015. Total number of catches was 65 and 33 in 2014 and 2015, respectively. Ethanol or ethanol + monochamol (a base blend) were not attractive to M. saltuarius with no difference from the control. Addition of α‐pinene and ipsdienol to the base blend did not significantly increase catches. However, ipsenol was significantly synergistic to the base blend in attracting M. saltuarius in 2014, and the blend (ipsenol + base blend) attracted meaningfully higher numbers of M. saltuarius in 2015. Our study illustrates the potential for monochamol and ipsenol baits for monitoring and trapping of M. saltuarius in the field.     

Evaluation of attractants and traps for monitoring small banded pine weevil Pissodes castaneus

This study aimed to develop a semiochemical‐baited trapping system to monitor the populations of small banded pine weevil, Pissodes castaneus, a serious pest in Pinus sylvestris young stands that are weakened by biotic and abiotic factors. The scope of the work included the development of a dispenser for compounds (ethanol and α‐pinene) emitted by P. sylvestris and the pheromones of P. castaneus: grandisol and grandisal. Additionally, the effectiveness of beetle catches in different types of traps (unitrap, cross‐unitrap and long and short pipe traps) baited with a dispenser was assessed. The olfactometric studies showed that most of the newly hatched beetles that had not fed were attracted by a mixture of grandisol and grandisal. However, in the group of feeding beetles, half were attracted by a mixture of ethanol and α‐pinene. These results indicated that both pheromones and α‐pinene plus ethanol should be useful for capturing P. castaneus beetles. In the field trials, the highest efficiency was found in baited unitraps that caught up to several hundred P. castaneus beetles, while the baited cross‐unitraps caught up to a few dozen beetles. No insects were found in either type of baited pipe trap or in any of the unbaited control traps. The baited unitraps and cross‐unitraps also collected, with varied intensity, Hylobius abietis beetles, a serious pest of reforestations. These results indicate the possibility of using a unitrap baited with a 4‐component attractant for monitoring P. castaneus in integrated pest management for the protection of young forests.     

Ophiostomatoid fungi (Ascomycota) associated with <Emphasis Type="Italic">Pinus tabuliformis</Emphasis> infested by <Emphasis Type="Italic">Dendroctonus valens</Emphasis> (Coleoptera) in northern China and an assessment of their pathogenicity on mature trees

Dendroctonus valens is an invasive pest in coniferous forests of northern China. It was suspected of being responsible for the death of more than three million Pinus tabuliformis trees. The present study sought to identify the ophiostomatoid fungi associated with D. valens in northern China and understand the possible role of these fungi in the pine decline. On the basis of morphology, physiology, mating compatibility and phylogenetic analyses of multiple DNA sequences, seven species of ophiostomatoid fungi were isolated from and around D. valens galleries: Leptographium alethinum, Grosmannia koreana (teleomorph of L. koreanum), L. procerum, L. sinoprocerum, L. truncatum, Pesotum aureum and P. pini. All have been recorded for the first time in China. Among them, the occurrence of the dominant species L. procerum is positively linked to attack intensities of D. valens. The pathogenicity of four species (L. koreanum, L. procerum, L. sinoprocerum and L. truncatum) was tested on mature P. tabuliformis trees by stem inoculation. All inoculated strains caused significant necrotic lesions on the inner bark. However, L. koreanum and L. truncatum induced more extensive lesions than L. procerum and L. sinoprocerum. Their association with D. valens and the P. tabuliformis decline is discussed.     

Development of an improved attractive lure for the pine shoot beetle, Tomicus piniperda (Coleoptera: Scolytidae)

Abstract 1 The pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), is an exotic pest of pine, Pinus spp., and was first discovered in North America in 1992. 2 Although primary attraction to host volatiles has been clearly demonstrated for T. piniperda, the existence and role of secondary attraction to insect‐produced pheromones have been widely debated. 3 Currently, commercial lures for T. piniperda include only the host volatiles α‐pinene in North America and α‐pinene, terpinolene and (+)‐3‐carene in Europe. Several potential pheromone candidates have been identified for T. piniperda. 4 We tested various combinations of host volatiles and pheromone candidates in Michigan, U.S.A., and Ontario, Canada, to determine an optimal blend. 5 Attraction of T. piniperda was significantly increased when trans‐verbenol (95% pure, 3.2%cis‐verbenol content) was added with or without myrtenol to α‐pinene or to blends of α‐pinene and other kairomones and pheromone candidates. 6 Our results, together with other research demonstrating that trans‐verbenol is produced by T. piniperda, support the designation of trans‐verbenol as a pheromone for T. piniperda. A simple operational lure consisting of α‐pinene and trans‐verbenol is recommended for optimal attraction of T. piniperda.     

Field response of non‐target beetles to Ips sexdentatus aggregation pheromone and pine volatiles

Although mass trapping cannot be a definitive control measure, it is one of the few ones available to contain the destruction of millions of cubic metres of conifer forests perpetrated every year worldwide by bark beetles. However, using bark beetle aggregation pheromones during both monitoring and control programs may negatively affect other saproxylic insects. The aim of this study was to describe the response of both Ips sexdentatus and its saproxylic beetle associates, especially predators, to traps baited with a commercial blend of I. sexdentatus aggregation pheromone. Furthermore, the usefulness of adding pine volatiles, such as (?)‐α‐pinene and ethanol, to the pheromone was discussed. The commercial blend proved to be attractive to I. sexdentatus adults, both when used alone and together with pine volatiles. Pheromone attractiveness, however, was lessened by the addition of the volatiles. The pheromone blend proved to be attractive to Thanasimus formicarius, as well as to other predator species. Overall, although during our study, traps baited only with (?)‐α‐pinene and ethanol attracted some predator specimens, I. sexdentatus pheromone traps were more attractive. Our study confirms that calendar differences in flight activity between the bark beetle and its predators are substantial; therefore, they should be taken into account when planning control measures. According to our data, the commercial blend of I. sexdentatus pheromone seems to be the most effective, among the baits used, in catching I. sexdentatus adults, while reducing the impact on T. formicarius.     

Log colonization by Ips sexdentatus prevented by increasing host unsuitability signaled by verbenone

Sudden increments of breeding material after windstorms, forest fires, or inappropriate management practices help bark beetles such as Ips sexdentatus Boerner (Coleoptera: Curculionidae: Scolytinae) increase in numbers and colonize standing healthy pine trees. Preventing bark beetles from arriving to susceptible trees or logs may have great relevance for bark beetle management. Recent studies have reported inhibition of the aggregation response of I. sexdentatus using verbenone. Two field experiments were conducted to examine the effect of verbenone on the colonization pattern of this beetle. The first experiment tested the combined effect of trans‐conophthorin, a non‐host bark volatile with known repellent effect, and verbenone on Pinus sylvestris L. (Pinaceae) log piles of two sizes, but failed to protect them against I. sexdentatus attack when these two infochemicals were released at low rates. The results of this experiment suggested an interaction with the associated secondary bark beetle Orthotomicus erosus (Wollaston). A second experiment examined the response of I. sexdentatus and O. erosus to log piles that released verbenone at 0, 2, 10, or 40 mg day^?1. Although I. sexdentatus colonization of Pinus nigra Arnold logs was completely prevented at 40 mg day^?1, O. erosus could be found at all tested verbenone release rates. Besides verbenone, O. erosus colonization density and the height from which logs originated were the variables that best explained I. sexdentatus log colonization pattern. In addition, I. sexdentatus and O. erosus were rarely recorded colonizing the same log, and niche breadth analyses suggested that they excluded each other. The role of verbenone in the colonization process and its potential use in the prevention of population buildups of damaging bark beetles such as I. sexdentatus are discussed.     

Behaviours of phoretic mites (Acari) associated with Ips pini and Ips grandicollis (Coleoptera: Curculionidae) during host‐tree colonization

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Effect of associated fungi on the immunocompetence of red turpentine beetle larvae,Dendroctonus valens (Coleoptera: Curculionidae: Scolytinae)

Abstract Dendroctonus–fungus symbioses are often considered as the ideal model systems to study the development and maintenance of ectosymbioses, and diverse interactions, including antagonism, commensalism and mutualism, have been documented between these organisms. The red turpentine beetle, Dendroctonus valens LeConte (Coleoptera: Curculionidae: Scolytinae) is a pine‐killing invasive beetle in northern China. Fungi species Ophiostoma minus, Leptographium sinoprocerum, L. terebrantis and L. procerum were associated with this bark beetle. Antagonistic interactions between D. valens and its associated fungi, such as O. minus and L. sinoprocerum, have been demonstrated, but the underlying causes of this phenomenon are unknown. Here, we first found the two tested fungi species retarded the net weight gain of D. valens larvae after completing 3‐day feeding on their media. Furthermore, we provide direct evidence indicating the effect of associated fungi on the immunocompetence of D. valens larvae to explain the documented antagonism. Our results showed that the activity of phenoloxidase and total phenoloxidase in D. valens larvae were significantly upregulated by two strains of associated fungi, O. minus and L. sinoprocerum as compared with the controls. The phenoloxidase ratio increased significantly in the larvae which had fed for 3 days on media inoculated with O. minus. Because insect immune defenses are costly to be deployed, these results could be explored as one of the underlying mechanisms of the documented antagonism.     

红脂大小蠹种群空间格局地统计学分析及抽样技术

红脂大小蠹是我国一重要林业外来入侵害虫,主要危害油松、白皮松和华山松,目前已扩散到山西、陕西、河北、河南、北京等地。为了研究其种群空间格局,用地统计学方法分析了受害程度不同的油松纯林和混交林林分内红脂大小蠹种群空间分布特性。结果表明:两种受害程度不同的油松纯林内红脂大小蠹的危害具有显著差异,轻度受害林分内有虫株率为4.9%,而重度受害林分则达到23.6%。全方向的变异函数曲线图分析得知:轻度受害纯林、重度受害纯林、混交林林分内红脂大小蠹种群的空间依赖范围分别为49.400、47.400\,73.820m,而局部空间连续性强度分别为0.824、0.582\,0.762。轻度受害纯林与重度受害纯林林分内红脂大小蠹种群均表现为较强的空间聚集性,而混交林受害林分内其种群则表现为随机分布。受害程度不同的纯林林分内红脂大小蠹种群空间格局具有一定的差异,而且纯林与混交林林分内其种群空间格局也有较大的差别。本文根据红脂大小蠹种群空间分布特性,对其种群空间抽样技术与传统抽样技术做了相应的比较与探讨。     

Saccharide-mediated antagonistic effects of bark beetle fungal associates on larvae

Bark beetles are among the most destructive of pine forest pests and they form close symbiotic relationships with ophiostomatoid fungi. Although some fungi are considered to be mutualistic symbionts of bark beetles with respect to the supply of nutrients, detrimental effects of fungal symbionts on larval growth have also been frequently reported. The mechanisms of such antagonistic effects are hypothesized to be a decrease in nutritional resources caused by competition for saccharides by the fungi. Here, we provide experimental evidence that three beetle-associated fungi modify the nutritional content of an artificial phloem diet, leading to a detrimental effect on the growth of Dendroctonus valens larvae. When larvae were fed a diet of pine phloem in agar medium colonized with any of these fungi, feeding activity was not affected but weight significantly decreased. Additional analysis showed that fungi depleted the fructose and glucose concentrations in the phloem media. Furthermore, these detrimental effects were neutralized by supplementing the media with fructose or glucose, suggesting that fungi may affect larval growth by modifying diet saccharide contents. These data indicate that fungus-induced nutritional changes in bark beetle diet can affect larval growth, and that the mechanism involves fungus-induced saccharide depletion from the larval diet.