Influences of the biophysical environment on blister rust and mountain pine beetle,and their interactions,in whitebark pine forests

Aim To understand how the biophysical environment influences patterns of infection by non‐native blister rust (caused by Cronartium ribicola) and mortality caused by native mountain pine beetles (Dendroctonus ponderosae) in whitebark pine (Pinus albicaulis) communities, to determine how these disturbances interact, and to gain insight into how climate change may influence these patterns in the future. Location High‐elevation forests in south‐west Montana, central Idaho, eastern and western Oregon, USA. Methods Stand inventory and dendroecological methods were used to assess stand structure and composition and to reconstruct forest history at sixty 0.1‐ha plots. Patterns of blister rust infection and mountain pine beetle‐caused mortality in whitebark pine trees were examined using nonparametric Kruskal–Wallis ANOVA, Mann–Whitney U‐tests, and Kolmogorov–Smirnov two‐sample tests. Stepwise regression was used to build models of blister rust infection and mountain pine beetle‐related mortality rates based on a suite of biophysical site variables. Results Occurrence of blister rust infections was significantly different among the mountain ranges, with a general gradient of decreasing blister rust occurrence from east to west. Evidence of mountain pine beetle‐caused mortality was identified on 83% of all dead whitebark pine trees and was relatively homogenous across the study area. Blister rust infected trees of all ages and sizes uniformly, while mountain pine beetles infested older, larger trees at all sites. Stepwise regressions explained 64% and 58% of the variance in blister rust infection and beetle‐caused mortality, respectively, indicating that these processes are strongly influenced by the biophysical environment. More open stand structures produced by beetle outbreaks may increase the exposure of surviving whitebark pine trees to blister rust infection. Main conclusions Variability in the patterns of blister rust infection and mountain pine beetle‐caused mortality elucidated the fundamental dynamics of these disturbance agents and suggests that the effects of climate change will be complex in whitebark pine communities and vary across the species’ range. Interactions between blister rust and beetle outbreaks may accelerate declines or facilitate the rise of rust resistance in whitebark pine depending on forest conditions at the time of the outbreak.     

Increasing water‐use efficiency and age‐specific growth responses of old‐growth ponderosa pine trees in the Northern Rockies

We examined radial growth responses of ponderosa pine (Pinus ponderosa var. ponderosa) between 1905–1954 and 1955–2004 to determine if the effects of increased intrinsic water‐use efficiencies (iWUE) caused by elevated atmospheric CO₂ concentrations were age‐specific. We collected 209 cores from five sites in the Northern Rockies and calculated iWUE using carbon isotope data from 1850 to 2004. Standardized radial growth responses were age dependent, with older trees exhibiting significantly higher values than younger trees during the later period at four sites and all sites combined. No significant differences in radial growth existed either for the individual sites or combined site during the earlier period. Increases in iWUE during 1955–2004 were 11% greater than during 1905–1954, and pentadal fluctuations in iWUE were significantly correlated with the radial growth of older trees from 1850 to 2004. Radial growth of younger trees and iWUE were not significantly correlated. Our results suggest that: (1) responses to elevated atmospheric CO₂ in old‐growth ponderosa forests are age‐specific; (2) radial growth increases in older trees coincided with increased iWUE; (3) ponderosa had increased growth rates in their third, fourth, and fifth centuries of life; and (4) age‐specific growth responses during 1955–2004 are unique since at least the mid‐16th century.     

Area‐wide application of verbenone‐releasing flakes reduces mortality of whitebark pine Pinus albicaulis caused by the mountain pine beetle Dendroctonus ponderosae

• 1 DISRUPT Micro‐Flake Verbenone Bark Beetle Anti‐Aggregant flakes (Hercon Environmental, Inc., Emigsville, Pennsylvania) were applied in two large‐scale tests to assess their efficacy for protecting whitebark pine Pinus albicaulis Engelm. from attack by mountain pine beetle Dendroctonus ponderosae Hopkins (Coleoptera: Scolytinae) (MPB). At two locations, five plots of equivalent size and stand structure served as untreated controls. All plots had early‐ to mid‐outbreak beetle populations (i.e. 7.1–29.2 attacked trees/ha). Verbenone was applied at 370 g/ha in both studies. Intercept traps baited with MPB aggregation pheromone were placed near the corners of each plot after the treatment in order to monitor beetle flight within the plots. Trap catches were collected at 7‐ to 14‐day intervals, and assessments were made at the end of the season of stand structure, stand composition and MPB attack rate for the current and previous years.
• 2 Applications of verbenone flakes significantly reduced the numbers of beetles trapped in treated plots compared with controls at both sites by approximately 50% at the first collection date.
• 3 The applications also significantly reduced the proportion of trees attacked in both Wyoming and Washington using the proportion of trees attacked the previous year as a covariate in the model for analysis of current year attack rates; in both sites, the reduction was ≥ 50%.
• 4 The flake formulation of verbenone appears to have promise for area‐wide treatment by aerial application when aiming to control the mountain pine beetle in whitebark pine forests.

     

Increased water‐use efficiency translates into contrasting growth patterns of Scots pine and sessile oak at their southern distribution limits

In forests, the increase in atmospheric CO₂ concentrations (C_a) has been related to enhanced tree growth and intrinsic water‐use efficiency (iWUE). However, in drought‐prone areas such as the Mediterranean Basin, it is not yet clear to what extent this “fertilizing” effect may compensate for drought‐induced growth reduction. We investigated tree growth and physiological responses at five Scots pine (Pinus sylvestris L.) and five sessile oak (Quercus petraea (Matt.) Liebl.) sites located at their southernmost distribution limits in Europe for the period 1960–2012 using annually resolved tree‐ring width and δ¹³C data to track ecophysiological processes. Results indicated that all 10 natural stands significantly increased their leaf intercellular CO₂ concentration (C_i), and consequently iWUE. Different trends in the theoretical gas‐exchange scenarios as a response to increasing C_a were found: generally, C_i tended to increase proportionally to C_a, except for trees at the driest sites in which C_i remained constant. C_i from the oak sites displaying higher water availability tended to increase at a comparable rate to C_a. Multiple linear models fitted at site level to predict basal area increment (BAI) using iWUE and climatic variables better explained tree growth in pines (31.9%–71.4%) than in oak stands (15.8%–46.8%). iWUE was negatively linked to pine growth, whereas its effect on growth of oak differed across sites. Tree growth in the western and central oak stands was negatively related to iWUE, whereas BAI from the easternmost stand was positively associated with iWUE. Thus, some Q. petraea stands might have partially benefited from the “fertilizing” effect of rising C_a, whereas P. sylvestris stands due to their strict closure of stomata did not profit from increased iWUE and consequently showed in general growth reductions across sites. Additionally, the inter‐annual variability of BAI and iWUE displayed a geographical polarity in the Mediterranean.     

Limited evidence for CO2‐related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO₂ may enhance photosynthesis and/or decrease stomatal conductance (g_s) thereby enhancing intrinsic water‐use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO₂ and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ¹³C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in g_s. Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing g_s may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.     

Predators and competitors of the mountain pine beetle Dendroctonus ponderosae (Coleoptera: Curculionidae) in stands of changing forest composition associated with elevation

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Woodpecker nest survival,density, and a pine beetle outbreak

Mountain pine beetle (Dendroctonus ponderosae) outbreaks in western North American coniferous forests are increasing in size and severity. An understanding of wildlife population responses to pine beetle outbreaks is needed to inform habitat conservation strategies. We monitored 355 nests of 5 woodpecker species during 2 sampling periods, before (2003–2006) and after (2009–2014) the peak of a pine beetle outbreak in dry mixed conifer forest of Montana, USA. Three of 5 woodpecker species represented the beetle-foraging group: American three-toed (Picoides dorsalis), hairy (Dryobates villosus), and downy (D. pubescens) woodpeckers. The other 2 species studied were northern flicker (Colaptes auratus), a foraging and habitat generalist, and red-naped sapsucker (Sphyrapicus nuchalis), a sap forager and bark-gleaning insectivore. We analyzed daily survival rate of nests in relation to pine beetle outbreak (445,000 ha) severity and timing, along with covariates unrelated to the outbreak (temp, nest height, and nest tree diameter). Our results provided stronger evidence for relationships between woodpecker nest survival and the non-outbreak variables than those associated with outbreaks. Our results indicated limited support for nest survival relationships with beetle severity (annual and cumulative pine tree mortality at 0.81-ha and 314-ha scales). Nevertheless, we observed a significant increase in densities of hatched nests for beetle-foraging woodpeckers following the outbreak. Our results suggest that woodpeckers, particularly beetle foragers, respond numerically to pine beetle outbreaks through increased nesting densities more so than functionally via nest survival. © 2019 The Authors. Journal of Wildlife Management Published by Wiley Periodicals, Inc. on behalf of The Wildlife Society.     

Tree‐mediated interactions between the jack pine budworm and a mountain pine beetle fungal associate

1. Coniferous trees deploy a combination of constitutive (pre‐existing) and induced (post‐invasion), structural and biochemical defences against invaders. Induced responses can also alter host suitability for other organisms sharing the same host, which may result in indirect, plant‐mediated interactions between different species of attacking organisms. 2. Current range and host expansion of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB) from lodgepole pine‐dominated forests to the jack pine‐dominated boreal forests provides a unique opportunity to investigate whether the colonisation of jack pine (Pinus banksiana Lamb.) by MPB will be affected by induced responses of jack pine to a native herbaceous insect species: the jack pine budworm (Choristoneura pinus pinus Freeman; JPBW). 3. We simulated MPB attacks with one of its fungal associates, Grosmannia clavigera Robinson‐Jeffrey & Davidson, and tested induction of either herbivory by JPBW or inoculation with the fungus followed by a challenge treatment with the other organism on jack pine seedlings and measured and compared monoterpene responses in needles. 4. There was clear evidence of an increase in jack pine resistance to G. clavigera with previous herbivory, indicated by smaller lesions in response to fungal inoculations. In contrast, although needle monoterpenes greatly increased after G. clavigera inoculation and continued to increase during the herbivory challenge, JPBW growth was not affected, but JPBW increased the feeding rate to possibly compensate for altered host quality. 5. Jack pine responses varied greatly and depended on whether seedlings were treated with single or multiple organisms, and their order of damage.     

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.     

Effects of pine wilt disease invasion on soil properties and Masson pine forest communities in the Three Gorges reservoir region,China

Pine wilt disease (PWD) has caused significant Masson pine mortality in the Three Gorges reservoir region in central China. In this study, five uniform Masson pine stand types infected by PWD were selected and surveyed on slopes and aspects with similar environmental conditions. In sites that had been infected, soil bulk density was reduced, and the difference among the groups was statistically significant (P < 0.05) at the 0–10 cm and 10–20 cm soil layers, but not at 20–40 cm. Other soil water‐related physical properties, excluding noncapillary porosity, significantly differed among the groups in all soil layers. Additionally, the values of available phosphorus, sodium, potassium, calcium, and magnesium were higher in the invaded stands, but the total nitrogen and organic matter contents were lower. Masson pine does not become reestablished following PWD‐induced mortality but is instead replaced by broad‐leaved tree species. Among the 19 examined environmental variables, five were found to be significantly related with the ordination of plant community structure: Masson pine stumps (MPS), K⁺, capillary water holding capacity (CWHC), capillary porosity (CP), and soil water content (SWC). Among these factors, the plant community structure was principally related to MPS and K⁺. The findings of this study show that the outbreak of PWD has impacted Masson pine forest soil properties and altered forest community composition. The disease is negatively related with the presence of Masson pine and positively associated with that of broad‐leaved tree species.     

Mountain pine beetle host-range expansion threatens the boreal forest

The current epidemic of the mountain pine beetle (MPB), an indigenous pest of western North American pine, has resulted in significant losses of lodgepole pine. The leading edge has reached Alberta where forest composition shifts from lodgepole to jack pine through a hybrid zone. The susceptibility of jack pine to MPB is a major concern, but there has been no evidence of host-range expansion, in part due to the difficulty in distinguishing the parentals and their hybrids. We tested the utility of a panel of microsatellite loci optimized for both species to classify lodgepole pine, jack pine and their hybrids using simulated data. We were able to accurately classify simulated individuals, and hence applied these markers to identify the ancestry of attacked trees. Here we show for the first time successful MPB attack in natural jack pine stands at the leading edge of the epidemic. This once unsuitable habitat is now a novel environment for MPB to exploit, a potential risk which could be exacerbated by further climate change. The consequences of host-range expansion for the vast boreal ecosystem could be significant.     

Attack dynamics of the pine shoot beetle, Tomicus piniperda (Col.; Scolytinae) in Scots pine stands defoliated by Bupalus piniaria (Lep.; Geometridae)

Abstract 1 After a 1‐year, extensive pine looper (Bupalus piniaria) outbreak, plots were laid out to study tree susceptibility to attack, and performance of Tomicus piniperda in pine trees suffering from varying levels of defoliation. 2 Tomicus piniperda was the dominating stem‐attacking species among the primary stem colonizers, and 82% of all trees that died had been colonized by T. piniperda. 3 Beetle attacks primarily struck severely defoliated trees, i.e. trees that suffered from 90% to 100% defoliation. 4 Beetle attacks peaked in the second year after cessation of the outbreak, and suppressed trees were both more frequently attacked and more susceptible to beetle attack than intermediate and dominant trees. 5 Trees surviving beetle attacks carried more foliage than trees that did not survive the attacks. 6 A single year of severe defoliation is enough to render pine trees susceptible to secondary pests, such as T. piniperda.     

Spatial–temporal analysis of species range expansion: the case of the mountain pine beetle, Dendroctonus ponderosae

Aim The spatial extent of western Canada’s current epidemic of mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is increasing. The roles of the various dispersal processes acting as drivers of range expansion are poorly understood for most species. The aim of this paper is to characterize the movement patterns of the mountain pine beetle in areas where range expansion is occurring, in order to describe the fine‐scale spatial dynamics of processes associated with mountain pine beetle range expansion. Location Three regions of Canada’s Rocky Mountains: Kicking Horse Pass, Yellowhead Pass and Pine Pass. Methods Data on locations of mountain pine beetle‐attacked trees of predominantly lodgepole pine (Pinus contorta var. latifolia) were obtained from annual fixed‐wing aircraft surveys of forest health and helicopter‐based GPS surveys of mountain pine beetle‐damaged areas in British Columbia and Alberta. The annual (1999–2005) spatial extents of outbreak ranges were delineated from these data. Spatial analysis was conducted using the spatial–temporal analysis of moving polygons (STAMP), a recently developed pattern‐based approach. Results We found that distant dispersal patterns (spot infestations) were most often associated with marginal increases in the areal size of mountain pine beetle range polygons. When the mountain pine beetle range size increased rapidly relative to the years examined, local dispersal patterns (adjacent infestation) were more common. In Pine Pass, long‐range dispersal (> 2 km) markedly extended the north‐east border of the mountain pine beetle range. In Yellowhead Pass and Kicking Horse Pass, the extension of the range occurred incrementally via ground‐based spread. Main conclusions Dispersal of mountain pine beetle varies with geography as well as with host and beetle population dynamics. Although colonization is mediated by habitat connectivity, during periods of low overall habitat expansion, dispersal to new distant locations is common, whereas during periods of rapid invasion, locally connected spread is the dominant mode of dispersal. The propensity for long‐range transport to establish new beetle populations, and thus to be considered a driver of range expansion, is likely to be determined by regional weather patterns, and influenced by local topography. We conclude that STAMP appears to be a useful approach for examining changes in biogeograpical ranges, with the potential to reveal both fine‐ and large‐scale patterns.     

Wood anatomy and carbon‐isotope discrimination support long‐term hydraulic deterioration as a major cause of drought‐induced dieback

Hydraulic impairment due to xylem embolism and carbon starvation are the two proposed mechanisms explaining drought‐induced forest dieback and tree death. Here, we evaluate the relative role played by these two mechanisms in the long‐term by quantifying wood‐anatomical traits (tracheid size and area of parenchyma rays) and estimating the intrinsic water‐use efficiency (iWUE) from carbon isotopic discrimination. We selected silver fir and Scots pine stands in NE Spain with ongoing dieback processes and compared trees showing contrasting vigour (declining vs nondeclining trees). In both species earlywood tracheids in declining trees showed smaller lumen area with thicker cell wall, inducing a lower theoretical hydraulic conductivity. Parenchyma ray area was similar between the two vigour classes. Wet spring and summer conditions promoted the formation of larger lumen areas, particularly in the case of nondeclining trees. Declining silver firs presented a lower iWUE than conspecific nondeclining trees, but the reverse pattern was observed in Scots pine. The described patterns in wood anatomical traits and iWUE are coherent with a long‐lasting deterioration of the hydraulic system in declining trees prior to their dieback. Retrospective quantifications of lumen area permit to forecast dieback in declining trees 2–5 decades before growth decline started. Wood anatomical traits provide a robust tool to reconstruct the long‐term capacity of trees to withstand drought‐induced dieback.     

Modeling mountain pine beetle (Dendroctonus ponderosae) oviposition

Mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae, Scolytinae), is a significant forest disturbance agent with a widespread distribution in western North America. Population success is influenced by temperatures that drive phenology and ultimately the adult emergence synchrony required to mass attack and kill host trees during outbreaks. In addition to lifestage‐specific developmental rates and thresholds, oviposition timing can be a source of variance in adult emergence synchrony, and is a critical aspect of mountain pine beetle phenology. Adaptation to local climates has resulted in longer generation times in southern compared to northern populations in common gardens, and the role of oviposition rate in these differences is unclear. Oviposition rates and fecundity in a northern population have been described, although data are lacking for southern populations. We assessed southern mountain pine beetle oviposition rates and fecundity in a range of temperatures using a non‐destructive technique that included frequent X‐ray imaging. We found that oviposition rate and fecundity vary independently such that a female with high oviposition rate did not necessarily have high fecundity and vice versa. Observed fecundity within the 30‐day experimental period was lowest at the lowest temperature, although estimated potential fecundity did not differ among temperatures. Females at varying temperatures have the potential to lay similar numbers of eggs, although it will take longer at lower temperatures. Southern mountain pine beetle reared in Pinus strobiformis Engelm. (Pinaceae) had a higher upper threshold for oviposition, a similar lower threshold, and slightly greater potential fecundity compared to a northern population reared in Pinus contorta Douglas. A comparison of modeled oviposition rates between the two populations, which could be influenced by host tree, suggests that differences in oviposition rate do not explain observed differences in total generation time. Our oviposition model will facilitate development of a phenology model for southern mountain pine beetle populations.     

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.     

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.     

Post-fire Aleppo pine growth,C and N isotope composition depend on site dryness

Key message

The post-fire growth responses and changes in wood C and N isotope composition depend on site water availability and fire severity in Mediterranean Aleppo pine forests.

Abstract

Mediterranean forests are subjected to recurrent wildfires and summer droughts. Under warmer and drier conditions, it is required to determine how Mediterranean pines recover after wildfires, and how this translates into changes in tree radial growth and function (e.g. intrinsic water-use efficiency—iWUE). We analysed four Aleppo pine areas located in SE Spain affected by 1994 wildfires and subjected to different water availability, ranging from mesic to semi-arid conditions. We combined dendrochronological analyses with δ¹³C and δ¹⁵N wood isotopes to quantify the changes in radial growth (expressed as Basal Area Increment—BAI) and functional responses (iWUE and N cycling) to three fire severities (unburned sites, low and medium severities). We expected that the post-fire release in nutrients and a reduction in competition for water would enhance radial growth. We found that fire did not significantly alter growth patterns at the driest sites, but increased BAI at the wettest sites. δ¹³C was significantly (P ≤ 0.01) more negative only in burned stands located at the wettest site indicating a decreased iWUE and thus improved water availability. However, the δ¹⁵N was higher in severely burned than in unburned plots from all sites but the wettest site, indicating a potential fertilization effect of fire in sites subjected to mild drought severity. Site water availability determined how fire affected subsequent modifications in growth and tree functioning of Aleppo pine forests, that is, changes in iWUE and N cycling. Therefore, site dryness should be explicitly considered to forecast the growth and functioning responses of Mediterranean pine forests to the predicted increasing recurrence of fire events due to global warming.

     

Dispersal capacity of Monochamus galloprovincialis,the European vector of the pine wood nematode,on flight mills

The pine wood nematode (Bursaphelenchus xylophilus), which causes the symptoms of pine wilt disease, is recognized worldwide as a major forest pest. It was introduced into Portugal in 1999. It is transmitted between trees almost exclusively by longhorn beetles of the genus Monochamus, including, in particular, M. galloprovincialis (Coleoptera: Cerambycidae) in maritime pine forests. Accurate estimates of the flight capacity of this insect vector are required if we are to understand and predict the spread of pine wilt disease in Europe. Using computer‐linked flight mills, we evaluated the distance flown, the flight probability and speed of M. galloprovincialis throughout adulthood and investigated the effects of age, sex and body weight on these flight performances, which are proxies for dispersal capacity. The within‐population variability of flight performance in M. galloprovincialis was high, with a mean distance of 16 km flown over the lifetime of the beetle. Age and body weight had a significant positive effect on flight capacity, but there was no difference in performance between males and females. These findings have important implications for managing the spread of the pine wood nematode in European forests.     

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.