Driving factors of a vegetation shift from Scots pine to pubescent oak in dry Alpine forests

An increasing number of studies have reported on forest declines and vegetation shifts triggered by drought. In the Swiss Rhone valley (Valais), one of the driest inner‐Alpine regions, the species composition in low elevation forests is changing: The sub‐boreal Scots pine (Pinus sylvestris L.) dominating the dry forests is showing high mortality rates. Concurrently the sub‐Mediterranean pubescent oak (Quercus pubescens Willd.) has locally increased in abundance. However, it remains unclear whether this local change in species composition is part of a larger‐scale vegetation shift. To study variability in mortality and regeneration in these dry forests we analysed data from the Swiss national forest inventory (NFI) on a regular grid between 1983 and 2003, and combined it with annual mortality data from a monitoring site. Pine mortality was found to be highest at low elevation (below 1000 m a.s.l.). Annual variation in pine mortality was correlated with a drought index computed for the summer months prior to observed tree death. A generalized linear mixed‐effects model indicated for the NFI data increased pine mortality on dryer sites with high stand competition, particularly for small‐diameter trees. Pine regeneration was low in comparison to its occurrence in the overstorey, whereas oak regeneration was comparably abundant. Although both species regenerated well at dry sites, pine regeneration was favoured at cooler sites at higher altitude and oak regeneration was more frequent at warmer sites, indicating a higher adaptation potential of oaks under future warming. Our results thus suggest that an extended shift in species composition is actually occurring in the pine forests in the Valais. The main driving factors are found to be climatic variability, particularly drought, and variability in stand structure and topography. Thus, pine forests at low elevations are developing into oak forests with unknown consequences for these ecosystems and their goods and services.     

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.     

Novel forest decline triggered by multiple interactions among climate,an introduced pathogen and bark beetles

Novel forest decline is increasing due to global environmental change, yet the causal factors and their interactions remain poorly understood. Using tree ring analyses, we show how climate and multiple biotic factors caused the decline of whitebark pine (Pinus albicaulis) in 16 stands in the southern Canadian Rockies. In our study area, 72% of whitebark pines were dead and 18% had partially dead crowns. Tree mortality peaked in the 1970s; however, the annual basal area increment of disturbed trees began to decline significantly in the late 1940s. Growth decline persisted up to 30 years before trees died from mountain pine beetle (Dendroctonus ponderosae), Ips spp. bark beetles or non‐native blister rust pathogen (Cronartium ribicola). Climate–growth relations varied over time and differed among the healthy and disturbed subpopulations of whitebark pine. Prior to the 1940s, cool temperatures limited the growth of all subpopulations. Growth of live, healthy trees became limited by drought during the cool phase (1947 –1976) of the Pacific Decadal Oscillation (PDO) and then reverted to positive correlations with temperature during the subsequent warm PDO phase. In the 1940s, the climate–growth relations of the disturbed subpopulations diverged from the live, healthy trees with trees ultimately killed by mountain pine beetle diverging the most. We propose that multiple factors interacted over several decades to cause unprecedented rates of whitebark pine mortality. Climatic variation during the cool PDO phase caused drought stress that may have predisposed trees to blister rust. Subsequent decline in snowpack and warming temperatures likely incited further climatic stress and with blister rust reduced tree resistance to bark beetles. Ultimately, bark beetles and blister rust contributed to tree death. Our findings suggest the complexity of whitebark pine decline and the importance of considering multiway drought–disease–insect interactions over various timescales when interpreting forest decline.     

Drought as an Inciting Mortality Factor in Scots Pine Stands of the Valais, Switzerland

During the 20th century, high mortality rates of Scots pine (Pinus silvestris L.) have been observed over large areas in the Rhône valley (Valais, Switzerland) and in other dry valleys of the European Alps. In this study, we evaluated drought as a possible inciting factor of Scots pine decline in the Valais. Averaged tree-ring widths, standardized tree-ring series, and estimated annual mortality risks were related to a drought index. Correlations between drought indices and standardized tree-ring series from 11 sites showed a moderate association. Several drought years and drought periods could be detected since 1864 that coincided with decreased growth. Although single, extreme drought years had generally a short-term, reversible effect on tree growth, multi-year drought initiated prolonged growth decreases that increased a tree’s long-term risk of death. Tree death occurred generally several years or even decades after the drought. In conclusion, drought has a limiting effect on tree growth and acts as a bottleneck event in triggering Scots pine decline in the Valais.     

Factors influencing bark beetle outbreaks after forest fires on the Iberian Peninsula

Fires are among the most globally important disturbances in forest ecosystems. Forest fires can be followed by bark beetle outbreaks. Therefore, the dynamic interactions between bark beetle outbreaks and fire appear to be of general importance in coniferous forests throughout the world. We tested three hypotheses of how forest fires in pine ecosystems (Pinus pinaster Alton and P. radiata D. Don) in Spain could alter the population dynamics of bark beetles and influence the probability of further disturbance from beetle outbreaks: fire could affect the antiherbivore resin defenses of trees, change their nutritional suitability, or affect top-down controls on herbivore populations. P. radiata defenses decreased immediately after fire, but trees with little crown damage soon recovered with defenses higher than before. Fire either reduced or did not affect nutritional quality of phloem and either reduced or had no effect on the abundance, diversity, and relative biomass of natural enemies. After fire, bark beetle abundance increased via rapid aggregation of reproductive adults on scorched trees. However, our results indicate that for populations to increase to an outbreak situation, colonizing beetles must initiate attacks before tree resin defenses recover, host trees must retain enough undamaged phloem to facilitate larval development, and natural enemies should be sufficiently rare to permit high beetle recruitment into the next generation. Coincidence of these circumstances may promote the possibility of beetle populations escaping to outbreak levels.     

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.     

Observations and modeling of aboveground tree carbon stocks and fluxes following a bark beetle outbreak in the western United States

Bark beetle epidemics result in tree mortality across millions of hectares in North America. However, few studies have quantified impacts on carbon (C) cycling. In this study, we quantified the immediate response and subsequent trajectories of stand‐level aboveground tree C stocks and fluxes using field measurements and modeling for a location in central Idaho, USA that experienced an outbreak of mountain pine beetle (Dendroctonus ponderosae Hopkins). We measured tree characteristics in lodgepole pine (Pinus contorta) plots spanning a range of structure and mortality conditions. We then initialized the forest vegetation simulator, an individual tree‐based model, with these measurements and simulated the response of aboveground production of C fluxes as well as trajectories of C stocks and fluxes in the coming decades. Mountain pine beetles killed up to 52% of the trees within plots, with more larger trees killed. C stocks in lodgepole pine were reduced by 31–83% following the outbreak, and plot‐level C fluxes decreased 28–73%. Modeled C stocks increased nearly continuously following the infestation, recovering to preoutbreak levels in 25 years or less. Simulated aboveground tree C fluxes increased following the immediate postoutbreak decrease, then subsequently declined. Substantial variability of C stocks and fluxes among plots resulted from the number and size of killed and surviving trees. Our study illustrates that bark beetle epidemics alter forest C cycling unlike stand‐replacement wildfires or clear‐cut harvests, due in part to incomplete mortality coupled with the preference by beetles for larger trees. The dependency of postoutbreak C stocks and fluxes on stand structure suggests that C budget models and studies in areas experiencing mountain pine beetle disturbances need to include size distribution of trees for the most accurate results.     

Induced defence reaction in Scots pine following stem attacks by Tomicus piniperda

The induced defence reaction of Scots pine Pinus sylvestris to stem attacks by the common pine shoot beetle, Tomicus piniperda (Col, Scolytidae), was studied by inducing natural attacks on young Scots pine trees of different vigour Pruned trees were more heavily attacked by T piniperda than unpruned ones Vigorous trees successfully contained the attacking beetles within resinous lesions, whereas less vigorous trees failed to defend themselves The content of resin acids increased dramatically in the lesions on all trees, and the resin acid composition differed somewhat between some treatments Starch accumulation in the inner bark of the main stem was lower in pruned trees than in unpruned ones, and starch reserves were depleted in the most heavily attacked trees All but one of the unpruned trees survived the attack, whereas most of the severely pruned ones died following attack by beetles at densities exceeding c 300 egg galleries m⁻²     

Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Climate-driven tree growth and mortality in the Black Forest,Germany—Long-term observations

Episodic tree mortality can be caused by various reasons. This study describes climate-driven tree mortality and tree growth in the Black Forest mountain range in Germany. It is based on a 68-year consistent data series describing the annual mortality of all trees growing in a forest area of almost 250 thousand ha. The study excludes mortality caused by storm, snow and ice, and fire. The sequence of the remaining mortality, the so-called “desiccated trees,” is analyzed and compared with the sequence of the climatic water balance during the growing season and the annual radial growth of Norway spruce in the Black Forest. The annual radial growth series covers 121 years and the climatic water balance series 140 years. These unique time series enable a quantitative assessment of multidecadal drought and heat impacts on growth and mortality of forest trees on a regional spatial scale. Data compiled here suggest that the mortality of desiccated trees in the Black Forest during the last 68 years is driven by the climatic water balance. Decreasing climatic water balance coincided with an increase in tree mortality and growth decline. Consecutive hot and dry summers enhance mortality and growth decline as a consequence of drought legacies lasting several years. The sensitivity of tree growth and mortality to changes in the climatic water balance increases with the decreasing trend of the climatic water balance. The findings identify the climatic water balance as the main driver of mortality and growth variation during the 68-year observation period on a landscape-scale including a variety of different sites. They suggest that bark beetle population dynamics modify mortality rates. They as well provide evidence that the mortality during the last 140 years never was as high as in the most recent years.     

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     

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.     

Bark beetle and woodborer responses to stand thinning and prescribed fire in northeastern US coastal and inland pitch pine barrens

1. Southern pine beetle, Dendroctonus frontalis, has expanded its range further into the northeastern United States. This expansion threatens rare and ecologically valuable interior and coastal pitch pine barrens.
2. Pitch pine barrens restoration and southern pine beetle infestation suppression often involve leaving downed dead wood that saproxylic insects can exploit.
3. Semiochemical-baited traps were used to investigate the response of bark beetles and woodborers to restoration treatments at Rocky Point State Forest and the Albany Pine Bush Preserve, examples of coastal and interior pitch pine forests, respectively.
4. A total of 29,598 saproxylic insects from 116 species of bark beetles and woodborers were captured at Rocky Point State Forest, while 23,117 individuals from 67 species were captured at Albany Pine Bush Preserve.
5. Ips spp. were abundant at both sites with 28%–47% and 42%–74% of total collections at Rocky Point State Forest and Albany Pine Bush Preserve, respectively.
6. Ips grandicollis did not respond to treatments at either site. However, Ips pini was found in higher numbers in thinned blocks in Rocky Point State Forest.

     

Importance of resin ducts in reducing ponderosa pine mortality from bark beetle attack

The relative importance of growth and defense to tree mortality during drought and bark beetle attacks is poorly understood. We addressed this issue by comparing growth and defense characteristics between 25 pairs of ponderosa pine (Pinus ponderosa) trees that survived and trees that died from drought-associated bark beetle attacks in forests of northern Arizona, USA. The three major findings of our research were: (1) xylem resin ducts in live trees were >10% larger (diameter), >25% denser (no. of resin ducts mm⁻²), and composed >50% more area per unit ring growth than dead trees; (2) measures of defense, such as resin duct production (no. of resin ducts year⁻¹) and the proportion of xylem ring area to resin ducts, not growth, were the best model parameters of ponderosa pine mortality; and (3) most correlations between annual variation in growth and resin duct characteristics were positive suggesting that conditions conducive to growth also increase resin duct production. Our results suggest that trees that survive drought and subsequent bark beetle attacks invest more carbon in resin defense than trees that die, and that carbon allocation to resin ducts is a more important determinant of tree mortality than allocation to radial growth.     

ALLOZYME AND MORPHOLOGICAL DIFFERENTIATION OF MOUNTAIN PINE BEETLES DENDROCTONUS PONDEROSAE HOPKINS (COLEOPTERA: SCOLYTIDAE) ASSOCIATED WITH HOST TREE

The purpose of this study was to determine whether mountain pine beetles utilizing different host species were differentiated for either morphological or protein variation. Genetic differentiation among host species has been reported for the southern pine beetle, the Douglas-fir beetle, the jeffrey pine beetle, and the mountain pine beetle. However, in these studies, the host trees were sampled at separate sites, and hence geographic variation and variation due to host tree were confounded. The mountain pine beetle occasionally utilizes three host trees (ponderosa pine, lodgepole pine, and limber pine) at single sites in Colorado. Five polymorphic enzyme loci and six morphological characters were used to describe beetles resident in different hosts. Differentiation within a site among host trees was detected at two of five polymorphic proteins, and for both size and morphological shape. The magnitude of genetic differentiation among hosts within a site was approximately equivalent to the magnitude of differentiation among sites. These data suggest that the species of host tree may be an important biotic factor associated with the genetic structure of bark beetle communities. The results are discussed in terms of their potential role in the process of speciation by host race formation.     

Survey and phylogenetic analysis of culturable microbes in the oral secretions of three bark beetle species

In a recent study, we reported a previously undescribed behavior in which a bark beetle exuded oral secretions containing bacteria that have antifungal properties, and hence defend their galleries against pervasive antagonistic Hyphomycete fungi. Actinobacteria, a group known for their antibiotic properties, were the most effective against fungi that invade the spruce beetle galleries. In the present study, we describe the isolation and identification of microorganisms from oral secretions of three bark beetles (Coleoptera: Curculionidae: Scolytinae): the spruce beetle, Dendroctonus rufipennis Kirby, the mountain pine beetle, Dendroctonus ponderosae Hopkins, and the pine engraver, Ips pini Say. Bacteria isolated from these three species span the major bacterial classes α-, β-, and γ-Proteobacteria, Firmicutes, Bacteroidetes, and Actinobacteria, except for D. ponderosae , which yielded no α-proteobacteria or Bacteroidetes isolates. Spruce beetles and pine engraver beetles had similar numbers of α-proteobacteria isolates, but pine engravers yielded twice as many Bacteroidetes isolates as spruce beetles. In contrast, mountain pine beetles yielded more isolates in the β- and γ-proteobacteria than spruce beetles and pine engravers. The highest percentage of Actinobacteria was obtained from spruce beetles, followed by pine engravers and mountain pine beetles. All of the fungal isolates obtained from the three beetle species were Ascomycetes. The greatest fungal diversity was obtained in spruce beetles, which had nine species, followed by pine engravers with five, and mountain pine beetles with one.     

Comparative study of the analytical hierarchy process,frequency ratio,and logistic regression models for predicting the susceptibility to Ips sexdentatus in Crimean pine forests

The six-toothed bark beetle Ips sexdentatus is one of the most important pests of coniferous trees that can cause extensive tree mortality, and change the structure and composition of forest ecosystems. Many abiotic and biotic factors affect the infestation of bark beetles. Early detection of forest stands predisposed to bark beetle infestations will benefit from reducing the impacts of possible infestations. The study focused on the production and comparison of Ips sexdentatus susceptibility maps using the analytical hierarchy process (AHP), frequency ratio (FR), and logistical regression (LR) models. The research was carried out in the Crimean pine forests of the Taşköprü Forest Enterprise in Kastamonu City in the Western Black Sea region of Türkiye. The eight main criteria used to produce the map were the stand structure, site index, crown closure, stand age, slope, elevation, maximum temperature, and solar radiation. The map of the infested stands was used for the models' validation. Crown closure was determined as the one of the most important factors in all three models. The receiver operating characteristic (ROC) curves and area under the curve (AUC) were used to determine the accuracy of the maps. The validation results showed that the AUC for the FR model was 0.747, for the AHP model was 0.716, and for the LR model was 0.638. The results revealed that the FR model was more accurate than the other models in producing an I. sexdentatus susceptibility map. Besides, the AHP model was also reasonably accurate. This study could help decision makers to produce bark beetle susceptibility maps easily and rapidly so they can take the necessary precautions to slow or prevent infestations.     

The upward shift in altitude of pine mistletoe (Viscum album ssp. austriacum) in Switzerland—the result of climate warming?

Pine mistletoe (Viscum album ssp. austriacum) is common in natural Scots pine (Pinus sylvestris L.) forests in the alpine Rhone Valley, Switzerland. This semi-parasite, which is regarded as an indicator species for temperature, increases the drought stress on trees and may contribute to the observed pine decline in the region. We recorded mistletoes on representative plots of the Swiss National Forest Inventory ranging from 450 to 1,550 m a.s.l. We found mistletoe on 37% of the trees and on 56% of all plots. Trees infested with mistletoe had a significantly higher mortality rate than non-infested trees. We compared the current mistletoe occurrence with records from a survey in 1910. The current upper limit, 1,250 m, is roughly 200 m above the limit of 1,000–1,100 m found in the earlier survey 100 years ago. Applying a spatial model to meteorological data we obtained monthly mean temperatures for all sites. In a logistic regression mean winter temperature, pine proportion and geographic exposition significantly explained mistletoe occurrence. Using mean monthly January and July temperatures for 1961–1990, we calculated Skres plant respiration equivalent (RE) and regressed it against elevation to obtain the RE value at the current mistletoe elevation limit. We used this RE value and temperature from 1870–1899 in the regression and found the past elevation limit to be at 1,060 m, agreeing with the 1910 survey. For the predicted temperature rise by 2030, the limit for mistletoe would increase above 1,600 m altitude.     

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.     

Drought and stand susceptibility to attacks by the European spruce bark beetle: A remote sensing approach

1. Several time-series analyses have demonstrated that after extreme summer drought bark beetle damage increased. However, studies predicting stand susceptibility over large spatial extents are limited by technical constraints in obtaining detailed, spatially-explicit data on infestation spot occurrence.
2. Using a unique dataset of georeferenced bark beetle infestation data, we tested whether the spatial variation of local growing conditions of forest stands, topography, and landscape variables modified the local occurrence of Ips typographus infestations after a severe hot drought in Central Europe.
3. Bark beetle infestation occurrence depended on soil-related aridity intensity, elevation, slope, and soil conditions. We showed that elevation interacted with growing conditions and topography. At low elevations, spruce forests growing on flat areas and wetter soils were more sensitive to the infestations. On the contrary, forests on steep slopes and soils with low water availability were rarely attacked. At the landscape scale, bark beetle damage increased with host tree cover but decreased with compositional diversity.
4. Our findings are generally consistent with the growth-differentiation balance hypothesis that predicts that trees growing under chronic dry conditions tend to be more resistant against biotic disturbances.
5. Spruce stands at low elevations located in homogeneous landscapes dominated by spruce were those more exposed to bark beetles in the initial phase of a drought-induced outbreak.