Wildfire provides refuge from local extinction but is an unlikely driver of outbreaks by mountain pine beetle

Bark beetle outbreaks and wildfire are important disturbances in conifer ecosystems, yet their interactions are not well understood. We evaluated whether fire injury increased susceptibility of lodgepole pines (Pinus contorta) to mountain pine beetle (Dendroctonus ponderosae Hopkins), how it influenced beetle reproductive success, and whether beetle population phase altered this interaction. Eight sites that experienced wildfire and eight unburned sites were examined in the Greater Yellowstone Ecosystem (USA). Half were in areas where D. ponderosae was undergoing outbreaks, and half were in areas with low populations. We examined 2056 trees one year after fire for burn injury and beetle attack. We quantified beetle reproductive success in a random sample of 106 trees, and measured gallery areas of D. ponderosae and competing subcortical herbivores in 79 additional trees. Baited flight traps sampled stand-level populations of subcortical herbivores and predators. Wildfire predisposed trees to D. ponderosae attack, but nonlinearly, with moderately injured trees being most preferred. This tree-level interaction was influenced by stand-level beetle population size, in that both healthy and fire-injured trees of all classes were attacked where populations were high, but no healthy trees, and only low and moderately injured trees were killed where populations were low. The number of adult brood produced per female was likewise curvilinear, being highest in moderately injured trees. This reflected an apparent trade-off, with high intraspecific competition arising from the large number of beetles needed to overcome defenses in healthy trees, vs. high interspecific competition and low substrate quality in more injured trees. These results suggest that fire-injured trees can provide a reservoir for D. ponderosae during periods when populations are too low to overcome defenses of healthy trees, and might otherwise face localized extinction. However, the likelihood of populations increasing from endemic to outbreak levels in response to increased susceptibility is offset by the opposing constraints of lower substrate quality and higher competitor load in severely injured hosts, and the relative scarcity of moderately injured trees. Wildfire may confer some reproductive increases to populations already outbreaking. We present a conceptual model of how these disturbances and inherent feedbacks interact to affect beetle population dynamics.     

Influence of global warming on forest coleopteran communities with special reference to ambrosia and bark beetles

Forest insect pests are one of the major disturbance factors in forest ecosystems and their outbreaks are expected to be more severe under the influence of global warming. Coleopterans are dominant among forest insects and their ecological functions include general detritivores, dead wood feeders, fungivores, herbivores, live wood feeders and predators. Ambrosia and bark beetles contribute to ecological succession of forests and, therefore, ecological functions of forests can be changed in response to their outbreaks. Mountain pine beetle (MPB) outbreaks are the most dramatic example of changes in the ecological functions of forest due to the outbreak of a forest insect pest altered by global warming. Composition of coleopteran species varies with latitude. However, composition of functional groups is consistent with latitude which indicates that resources available to beetles are consistent. In coleopteran communities, ambrosia and bark beetles can become dominant due to increases of dead or stressed trees due to the warming climate. This can also induce changes in the ecological functions of coleopterans, i.e. selective force to displace trees that have lower ecological fitness due to temperature increase. Therefore, recent increases in the density ambrosia and bark beetles offer a chance to study ecological processes in forests under the influence of global warming.     

Death of vigorous trees benefits bark beetles

Bark beetles (Coleoptera: Scolytidae) are commonly associated with live host trees that are stressed, a relationship that has been attributed to lower host defenses or greater nutritional quality of these trees. However, most bark beetle species commonly inhabit freshly dead trees where induced host defenses are absent. In this study, we investigate the role of tree vigor at the time of death for pine engraver bark beetles, Ips pini (Say), breeding in freshly dead jack pine, Pinus banksiana Lamb. As indices of tree vigor, we considered tree size, phloem thickness, and several measures of recent growth rate (last year's growth increment, mean annual increment and basal area increment in the past 5 and 10 years, and periodic growth ratio). We examined the relationship between these indices in three stands, aged 60, 77, and 126 years, and found that phloem thickness, previously shown to have a strong positive effect on bark beetle reproduction, was only weakly associated with tree growth rate and inconsistently related to tree size among the three stands. To examine the effects of tree vigor on pine engraver reproduction, we felled 20 trees of various sizes from the 77-year-old stand, and experimentally established breeding males and females in 25-cm-long sections. Offspring were collected and characteristics of breeding galleries were measured. Using stepwise regression, we consistently found that indices associated with tree growth rate best explained beetle reproductive performance, as they were positively related to parental male and female establishment on logs, female reproductive success, length of egg galleries, proportion of eggs resulting in emerged offspring, and negatively related to the length of the post-egg gallery. Surprisingly, phloem thickness had no unique effect on pine engraver reproduction, except for a weak negative effect on the establishment success of parental females. The strong effect of tree vigor observed in this study suggests that substantial mortality of vigorous trees, such as caused by windthrow, can contribute to significant increases in bark beetle populations that could trigger outbreaks in living trees. Received: 3 February 1999 / Accepted: 27 April 1999     

Dietary benefits of fungal associates to an eruptive herbivore: potential implications of multiple associates on host population dynamics

We used the mountain pine beetle (Dendroctonus ponderosae Hopkins) and its two fungal associates, Grosmannia clavigera and Ophiostoma montium, to study potential nutritional benefits of fungi to bark beetles. We tested for potential effects of feeding on phloem colonized by fungi on beetle performance in field and laboratory studies. The fungi increased nitrogen levels in the phloem of attacked trees by 40%, indicating that it may be an important source of dietary nitrogen for mountain pine beetles. However, nitrogen levels of phloem inoculated with fungi in the laboratory were similar to uncolonized phloem, indicating that the fungi may redistribute nitrogen from the sapwood to the phloem rather than increase absolute levels of nitrogen. Beetles emerging from attacked trees carrying G. clavigera were larger than beetles carrying O. montium, which in turn were larger than beetles lacking fungi. Results of experimental laboratory studies varied, likely because of differences in the growth and sporulation of fungi under artificial conditions. Results indicate that the two fungi may offer complementary benefits to the mountain pine beetle because larvae preferentially fed on phloem colonized by both fungi together over phloem colonized by one fungus or uncolonized phloem. Teneral adults preemergence fed on spores in pupal chambers when they were produced and consumed little phloem before emerging. Teneral adults mined extensively in the phloem before emerging when spores were not produced in the pupal chamber. Our results provide evidence for a nutritional role of fungi in the diet of bark beetles and show that multiple associates may differentially affect beetle performance, which could have important implications for bark beetle population dynamics.     

Interactions Among Fuel Management,Species Composition,Bark Beetles,and Climate Change and the Potential Effects on Forests of the Lake Tahoe Basin

Climate-driven increases in wildfires, drought conditions, and insect outbreaks are critical threats to forest carbon stores. In particular, bark beetles are important disturbance agents although their long-term interactions with future climate change are poorly understood. Droughts and the associated moisture deficit contribute to the onset of bark beetle outbreaks although outbreak extent and severity is dependent upon the density of host trees, wildfire, and forest management. Our objective was to estimate the effects of climate change and bark beetle outbreaks on ecosystem carbon dynamics over the next century in a western US forest. Specifically, we hypothesized that (a) bark beetle outbreaks under climate change would reduce net ecosystem carbon balance (NECB) and increase uncertainty and (b) these effects could be ameliorated by fuels management. We also examined the specific tree species dynamics—competition and release—that determined NECB response to bark beetle outbreaks. Our study area was the Lake Tahoe Basin (LTB), CA and NV, USA, an area of diverse forest types encompassing steep elevation and climatic gradients and representative of mixed-conifer forests throughout the western United States. We simulated climate change, bark beetles, wildfire, and fuels management using a landscape-scale stochastic model of disturbance and succession. We simulated the period 2010–2100 using downscaled climate projections. Recurring droughts generated conditions conducive to large-scale outbreaks; the resulting large and sustained outbreaks significantly increased the probability of LTB forests becoming C sources over decadal time scales, with slower-than-anticipated landscape-scale recovery. Tree species composition was substantially altered with a reduction in functional redundancy and productivity. Results indicate heightened uncertainty due to the synergistic influences of climate change and interacting disturbances. Our results further indicate that current fuel management practices will not be effective at reducing landscape-scale outbreak mortality. Our results provide critical insights into the interaction of drivers (bark beetles, wildfire, fuel management) that increase the risk of C loss and shifting community composition if bark beetle outbreaks become more frequent.     

Emergence patterns of univoltine and bivoltine Ips typographus (L.) populations and associated natural enemies

Control measures aiming at reducing bark beetle populations and preserving their natural enemies require a sound knowledge on their overwintering and emergence behaviour. These behavioural traits were investigated in univoltine and bivoltine populations of the European spruce bark beetle (Ips typographus [L.], Coleoptera: Scolytinae) and its predators and parasitoids over several consecutive years. In univoltine populations, roughly 50% of the bark beetles left their brood trees in fall together with most parasitoids and some significant predatory flies and beetles. In bivoltine populations, <10% of the second bark beetle generation emerged in fall and the remainder overwintered under the bark of their brood trees. Likewise, most predatory beetles and flies spent wintertime with their prey under the bark, while most parasitic wasps emerged in fall. The spring emergence of bivoltine predatory beetles was found to occur up to 3 weeks earlier than that of I. typographus, while that of the predatory flies and the parasitoids was delayed by up to 1 month. In univoltine populations, the bark beetles emerged several weeks prior to most antagonistic taxa. In the heat year 2003, three I. typographus generations were produced at the lower location, 36% of the third generation emerged in fall, while the proportions of overwintering predators remained largely the same as in previous years. Similar to their host, more parasitoids left their brood trees in fall after warm years. The results show that sanitation felling during winter probably kills most bark beetles in bivoltine populations, but also eliminates many natural enemies. In univoltine populations, sanitation felling might be less detrimental to both I. typographus and natural enemies because a fair fraction of their populations will already have left the trees before cutting. Warmer climates may affect the interactions of bark beetles and natural enemies and thus the impact of control measures.     

Host resistance elicited by methyl jasmonate reduces emission of aggregation pheromones by the spruce bark beetle, Ips typographus

We treated Norway spruce (Picea abies) stems with methyl jasmonate (MeJA) to determine possible quantitative and qualitative effects of induced tree defenses on pheromone emission by the spruce bark beetle Ips typographus. We measured the amounts of 2-methyl-3-buten-2-ol and (S)-cis-verbenol, the two main components of the beetle's aggregation pheromone, released from beetle entrance holes, along with phloem terpene content and beetle performance in MeJA-treated and untreated Norway spruce logs. As expected, phloem terpene levels were higher and beetle tunnel length was shorter (an indication of poor performance) in MeJA-treated logs relative to untreated logs. Parallel to the higher phloem terpene content and poorer beetle performance, beetles in MeJA-treated logs released significantly less 2-methyl-3-buten-2-ol and (S)-cis-verbenol, and the ratio between the two pheromone components was significantly altered. These results suggest that host resistance elicited by MeJA application reduces pheromone emission by I. typographus and alters the critical ratio between the two main pheromone components needed to elicit aggregation. The results also provide a mechanistic explanation for the reduced performance and attractivity observed in earlier studies when bark beetles colonize trees with elicited host defenses, and extend our understanding of the ecological functions of conifer resistance against bark beetles.     

Anatomical defences against bark beetles relate to degree of historical exposure between species and are allocated independently of chemical defences within trees

Conifers possess chemical and anatomical defences against tree‐killing bark beetles that feed in their phloem. Resins accumulating at attack sites can delay and entomb beetles while toxins reach lethal levels. Trees with high concentrations of metabolites active against bark beetle‐microbial complexes, and more extensive resin ducts, achieve greater survival. It is unknown if and how conifers integrate chemical and anatomical components of defence or how these capabilities vary with historical exposure. We compared linkages between phloem chemistry and tree ring anatomy of two mountain pine beetle hosts. Lodgepole pine, a mid‐elevation species, has had extensive, continual contact with this herbivore, whereas high‐elevation whitebark pines have historically had intermittent exposure that is increasing with warming climate. Lodgepole pine had more and larger resin ducts. In both species, anatomical defences were positively related to tree growth and nutrients. Within‐tree constitutive and induced concentrations of compounds bioactive against bark beetles and symbionts were largely unrelated to resin duct abundance and size. Fewer anatomical defences in the semi‐naïve compared with the continually exposed host concurs with directional differences in chemical defences. Partially uncoupling chemical and morphological antiherbivore traits may enable trees to confront beetles with more diverse defence permutations that interact to resist attack.     

Stand‐replacing fires reduce susceptibility of lodgepole pine to mountain pine beetle outbreaks in Colorado

Aim As climate change is increasing the frequency, severity and extent of wildfire and bark beetle outbreaks, it is important to understand how these disturbances interact to affect ecological patterns and processes, including susceptibility to subsequent disturbances. Stand‐replacing fires and outbreaks of mountain pine beetle (MPB), Dendroctonus ponderosae, are both important disturbances in the lodgepole pine, Pinus contorta, forests of the Rocky Mountains. In the current study we investigated how time since the last stand‐replacing fire affects the susceptibility of the stand to MPB outbreaks in these forests. We hypothesized that at a stand‐scale, young post‐fire stands (< c. 100–150 years old) are less susceptible to past and current MPB outbreaks than are older stands. Location Colorado, USA. Methods We used dendroecological methods to reconstruct stand‐origin dates and the history of outbreaks in 23 lodgepole pine stands. Results The relatively narrow range of establishment dates among the oldest trees in most sampled stands suggested that these stands originated after stand‐replacing or partially stand‐replacing fires over the past three centuries. Stands were affected by MPB outbreaks in the 1940s/1950s, 1980s and 2000s/2010s. Susceptibility to outbreaks generally increased with stand age (i.e. time since the last stand‐replacing fire). However, this reduced susceptibility of younger post‐fire stands was most pronounced for the 1940s/1950s outbreak, less so for the 1980s outbreak, and did not hold true for the 2000s/2010s outbreak. Main conclusions Younger stands may not have been less susceptible to the most recent outbreak because: (1) after stands reach a threshold age of > 100–150 years, stand age does not affect susceptibility to outbreaks, or (2) the high intensity of the most recent outbreak reduces the importance of pre‐disturbance conditions for susceptibility to disturbance. If the warm and dry conditions that contribute to MPB outbreaks concurrently increase the frequency and/or extent of severe fires, they may thereby mitigate the otherwise increased landscape‐scale susceptibility to outbreaks. Potential increases in severe fires driven by warm and dry climatic trends may lead to a negative feedback by making lodgepole pine stands less susceptible to future MPB outbreaks.     

A model for the conquest of a tree by bark beetles

A model for the interaction between attacking bark beetles and a tree is developed and discussed. It is shown that in addition to the more intuitive outcomes where the tree wins or the beetles win, it is also deduced that under certain conditions there may exist a stable coexistence (at least for some period of time) between the beetles and a living tree. Finally, it is demonstrated that the outcome of the tree-beetle interaction often depends on initial conditions such as the number of colonizing beetles.
These results are discussed with reference to empirical findings, as well as to the development of proper population dynamics models for bark beetles in a forest stand, and models developed for assisting forest managers in avoiding bark beetle outbreaks and for minimizing the damage caused by a bark beetle outbreak.     

Spruce beetle outbreak was not driven by drought stress: Evidence from a tree‐ring iso‐demographic approach indicates temperatures were more important

Climate change has amplified eruptive bark beetle outbreaks over recent decades, including spruce beetle (Dendroctonus rufipennis). However, for projecting future bark beetle dynamics there is a critical lack of evidence to differentiate how outbreaks have been promoted by direct effects of warmer temperatures on beetle life cycles versus indirect effects of drought on host susceptibility. To diagnose whether drought‐induced host‐weakening was important to beetle attack success we used an iso‐demographic approach in Engelmann spruce (Picea engelmannii) forests that experienced widespread mortality caused by spruce beetle outbreaks in the 1990s, during a prolonged drought across the central and southern Rocky Mountain region. We determined tree death date demography during this outbreak to differentiate early‐ and late‐dying trees in stands distributed across a landscape within this larger regional mortality event. To directly test for a role of drought stress during outbreak initiation we determined whether early‐dying trees had greater sensitivity of tree‐ring carbon isotope discrimination (?¹³C) to drought compared to late‐dying trees. Rather, evidence indicated the abundance and size of host trees may have modified ?¹³C responses to drought. ?¹³C sensitivity to drought did not differ among early‐ versus late‐dying trees, which runs contrary to previously proposed links between spruce beetle outbreaks and drought. Overall, our results provide strong support for the view that irruptive spruce beetle outbreaks across North America have primarily been driven by warming‐amplified beetle life cycles whereas drought‐weakened host defenses appear to have been a distant secondary driver of these major disturbance events.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

THE EVOLUTION OF AGRICULTURE IN BEETLES (CURCULIONIDAE: SCOLYTINAE AND PLATYPODINAE)

Abstract Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi—known as ambrosia fungi—that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free‐living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle‐fungus associations, enabling them to use the biomass‐rich resource that trees represent and set the stage for at least one origin of eusociality.     

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.     

A Decade of Streamwater Nitrogen and Forest Dynamics after a Mountain Pine Beetle Outbreak at the Fraser Experimental Forest,Colorado

Forests of western North America are currently experiencing extensive tree mortality from a variety of bark beetle species, and insect outbreaks are projected to increase under warmer, drier climates. Unlike the abrupt biogeochemical changes typical after wildfire and timber harvesting, the outcomes of insect outbreaks are poorly understood. The mountain pine bark beetle (Dendroctonus ponderosae) began to attack lodgepole pine (Pinus contorta) at the Fraser Experimental Forest in 2002 and spread throughout the research area by 2007. We compared streamwater nitrogen (N) from 2003 through 2012 with data from the previous two decades in four watersheds with distinct forest management histories, stand structures, and responses to the beetle outbreak. Watersheds dominated by old-growth had larger trees and lost 85% of overstory pine and 44% of total basal area to bark beetles. In contrast, managed watersheds containing a mixture of second-growth (30–60 year old) and old-growth (250- to 350-year old) had higher density of subcanopy trees, smaller mean tree diameter, and lower bark beetle-induced mortality (~26% of total basal area). Streamwater nitrate concentrations were significantly higher in old-growth watersheds during the outbreak than pre-outbreak levels during snowmelt and base flow seasons. In mixed-age stands, streamwater nitrate concentrations were unaffected by the outbreak. Beetle outbreak elevated inorganic N export 43 and 74% in two old-growth watersheds though the amounts of N released in streamwater were low (0.04 and 0.15 kg N ha^?1) relative to atmospheric inputs (<2% of annual N deposition). Increased height, diameter, and foliar N of measured in residual live trees augmented demand for N, far in excess of the change in N export during the outbreak. Reallocation of soil resources released after pine mortality to overstory and understory vegetation helps explain high nutrient retention in watersheds affected by bark beetle outbreaks.     

Evaluation of funnel traps for characterizing the bark beetle (Coleoptera: Scolytidae) communities in ponderosa pine forests of north-central Arizona

Lindgren funnel traps baited with aggregation pheromones are widely used to monitor and manage populations of economically important bark beetles (Coleoptera: Scolytidae). This study was designed to advance our understanding of how funnel trap catches assess bark beetle communities and relative abundance of individual species. In the second year (2005) of a 3-yr study of the bark beetle community structure in north-central Arizona pine (Pinus spp.) forests, we collected data on stand structure, site conditions, and local bark beetle-induced tree mortality at each trap site. We also collected samples of bark from infested (brood) trees near trap sites to identify and determine the population density of bark beetles that were attacking ponderosa pine, Pinus ponderosa Douglas ex Lawson, in the area surrounding the traps. Multiple regression models indicated that the number of Dendroctonus and Ips beetles captured in 2005 was inversely related to elevation of the trap site, and positively associated with the amount of ponderosa pine in the stand surrounding the site. Traps located closer to brood trees also captured more beetles. The relationship between trap catches and host tree mortality was weak and inconsistent in forest stands surrounding the funnel traps, suggesting that trap catches do not provide a good estimate of local beetle-induced tree mortality. However, pheromone-baited funnel trap data and data from gallery identification in bark samples produced statistically similar relative abundance profiles for the five species of bark beetles that we examined, indicating that funnel trap data provided a good assessment of species presence and relative abundance.     

Are bark beetle outbreaks less synchronous than forest Lepidoptera outbreaks?

Comparisons of intraspecific spatial synchrony across multiple epidemic insect species can be useful for generating hypotheses about major determinants of population patterns at larger scales. The present study compares patterns of spatial synchrony in outbreaks of six epidemic bark beetle species in North America and Europe. Spatial synchrony among populations of the Eurasian spruce bark beetle Ips typographus was significantly higher than for the other bark beetle species. The spatial synchrony observed in epidemic bark beetles was also compared with previously published patterns of synchrony in outbreaks of defoliating forest Lepidoptera, revealing a marked difference between these two major insect groups. The bark beetles exhibited a generally lower degree of spatial synchrony than the Lepidoptera, possibly because bark beetles are synchronized by different weather variables that are acting on a smaller scale than those affecting the Lepidoptera, or because inherent differences in their dynamics leads to more cyclic oscillations and more synchronous spatial dynamics in the Lepidoptera.     

The impact of phloem nutrients on overwintering mountain pine beetles and their fungal symbionts

In the low nutrient environment of conifer bark, subcortical beetles often carry symbiotic fungi that concentrate nutrients in host tissues. Although bark beetles are known to benefit from these symbioses, whether this is because they survive better in nutrient-rich phloem is unknown. After manipulating phloem nutrition by fertilizing lodgepole pine trees (Pinus contorta Douglas var. latifolia), we found bolts from fertilized trees to contain more living individuals, and especially more pupae and teneral adults than bolts from unfertilized trees at our southern site. At our northern site, we found that a larger proportion of mountain pine beetle (Dendroctonus ponderosae Hopkins) larvae built pupal chambers in bolts from fertilized trees than in bolts from unfertilized trees. The symbiotic fungi of the mountain pine beetle also responded to fertilization. Two mutualistic fungi of bark beetles, Grosmannia clavigera (Rob.-Jeffr. & R. W. Davidson) Zipfel, Z. W. de Beer, & M. J. Wingf. and Leptographium longiclavatum Lee, S., J. J. Kim, & C. Breuil, doubled the nitrogen concentrations near the point of infection in the phloem of fertilized trees. These fungi were less capable of concentrating nitrogen in unfertilized trees. Thus, the fungal symbionts of mountain pine beetle enhance phloem nutrition and likely mediate the beneficial effects of fertilization on the survival and development of mountain pine beetle larvae.     

The Red-belted Bracket (Fomitopsis pinicola) colonizes spruce trees early after bark beetle attack and persists

The Red-belted Bracket (Fomitopsis pinicola) is one of the major decomposers of coniferous wood in Europe and can reach high densities after outbreaks of bark beetles. However, factors of dead wood type and decay stage, which determine the growth of reproductive biomass, i.e. basidiomes, remain unclear. In 2013, we surveyed 1280 dead wood objects and vital trees in spruce stands killed by the bark beetle Ips typographus in 2012, 2002, 1992 and in undisturbed stands for the presence, number, mean basidiome size and total volume of basidiomes. Living basidiomes were equally abundant on dead wood 1, 11, and 21 y after bark beetle outbreak, but were lacking on living trees. Our results indicate that F. pinicola is an effective early colonizer of the huge resource pulse of dead wood caused by the outbreak of bark beetles and basidiomes can persist for 21 y.