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Conifers have evolved complex oleoresin terpene defenses against herbivores and pathogens. In co-evolved bark beetles, conifer terpenes also serve chemo-ecological functions as pheromone precursors, chemical barcodes for host identification, or nutrients for insect-associated microbiomes. We highlight the genomic, molecular and biochemical underpinnings of the large chemical space of conifer oleoresin terpenes and volatiles. Conifer terpenes are predominantly the products of the conifer terpene synthase (TPS) gene family. Terpene diversity is increased by cytochromes P450 of the CYP720B class. Many conifer TPS are multiproduct enzymes. Multisubstrate CYP720B enzymes catalyse multistep oxidations. We summarise known terpenoid gene functions in various different conifer species with reference to the annotated terpenoid gene space in a spruce genome. Overall, biosynthesis of terpene diversity in conifers is achieved through a system of biochemical radiation and metabolic grids. Expression of TPS and CYP720B genes can be specific to individual cell types of constitutive or traumatic resin duct systems. Induced terpenoid transcriptomes in resin duct cells lead to dynamic changes of terpene composition and quantity to fend off herbivores and pathogens. While terpenoid defenses have contributed much to the evolutionary success of conifers, under new conditions of climate change, these defences may become inconsequential against range-expanding forest pests.

Over the last decades, climate change has triggered an increase in the frequency of sprucebark beetle (Ips typographusL.) in Central Europe. More than 50% of forests in the Czech Republic areseriously threatened by this pest, leading to high ecological and economic losses. The exponentialincrease of bark beetle infestation hinders the implementation of costly field campaigns to prevent andmitigate its effects. Remote sensing may help to overcome such limitations as it provides frequent andspatially continuous data on vegetation condition. Using Sentinel-2 images as main input, two modelshave been developed to test the ability of this data source to map bark beetle damage and severity.All models were based on a change detection approach, and required the generation of previous forestmask and dominant species maps. The first damage mapping model was developed for 2019 and2020, and it was based on bi-temporal regressions in spruce areas to estimate forest vitality and barkbeetle damage. A second model was developed for 2020 considering all forest area, but excludingclear-cuts and completely dead areas, in order to map only changes in stands dominated by alivetrees. The three products were validated with in situ data. All the maps showed high accuracies (acc>0.80). Accuracy was higher than 0.95 and F1-score was higher than 0.88 for areas with high severity,with omission errors under 0.09 in all cases. This confirmed the ability of all the models to detectbark beetle attack at the last phases. Areas with no damage or low severity showed more complexresults. The no damage category yielded greater commission errors and relative bias (CEs=0.30-0.42,relB=0.42-0.51). The similar results obtained for 2020 leaving out clear-cuts and dead trees provedthat the proposed methods could be used to help forest managers fight bark beetle pests. These bioticdamage products based on Sentinel-2 can be set up for any location to derive regular forest vitalitymaps and inform of early damage.

1. During drought, the tree subpopulations (such as size or vigour classes) that suffer disproportionate mortality can be conceptually arrayed along a continuum defined by the actions of biotic agents, particularly insects. At one extreme, stress dominates: insects are absent or simply kill the most physiologically stressed trees. At the opposite extreme, host selection dominates: outbreaking insects kill trees independently of their stress, instead selecting trees based on size or other traits. Intermediate responses are also possible. Yet for mixed-species forests, we lack a broad understanding of the relative importance of insects in determining exactly which subpopulations of trees suffer disproportionate mortality during drought, and whether these subpopulations differ among co-occurring tree species. 2. During an extreme drought, we documented the roles of native bark beetles in the mortality of five tree species in California's Sierra Nevada. We analysed the patterns and agents of tree mortality in 12 permanent plots and the patterns of mortality in 89 temporary plots. 3. Most tree mortality was associated with bark beetles. However, the growth rates (an indicator of chronic stress) and sizes of trees that suffered greatest bark beetle-related mortality differed sharply among tree taxa, variously conforming with domination by stress (Abies concolor), domination by host selection (Pinus lambertiana and P. ponderosa) or a mix of the two (Calocedrus decurrens). Quercus kelloggii mortality remained relatively low. Thus, even during extreme drought substantial proportions of stressed trees survived because they were of sizes that mostly avoided fatal insect attack. Conversely, substantial proportions of comparatively unstressed trees died because they were of sizes that were selectively killed by outbreaking insects. 4. Synthesis. Native bark beetles were primarily responsible for determining which subpopulations of trees suffered greatest mortality during drought. However, idiosyncratic host-tree selection by the different bark beetle taxa meant that the tree subpopulations suffering greatest mortality differed strikingly among tree taxa—for example, high mortality of small trees of one species, but of large trees of another. If idiosyncratic host-tree selection by biotic mortality agents proves to be a generally common phenomenon, it could help explain weak broadscale correlations between tree traits and tree mortality during drought.

The bark beetle (Ips typographus) disturbance represents serious environmental and economic issue and presents a major challenge for forest management. A timely detection of bark beetle infestation is therefore necessary to reduce losses. Besides wood production, a bark beetle outbreak affects the forest ecosystem in many other ways including the water cycle, nutrient cycle, or carbon fixation. On that account, (not just) European temperate coniferous forests may become endangered ecosystems. Our study was performed in the unmanaged zone of the Krkonoše Mountains National Park in the northern part of the Czech Republic where the natural spreading of bark beetle is slow and, therefore, allow us to continuously monitor the infested trees that are, in contrast to managed forests, not being removed. The aim of this work is to evaluate possibilities of unmanned aerial vehicle (UAV)-mounted low-cost RGB and modified near-infrared sensors for detection of different stages of infested trees at the individual level, using a retrospective time series for recognition of still green but already infested trees (so-called green attack). A mosaic was created from the UAV imagery, radiometrically calibrated for surface reflectance, and five vegetation indices were calculated; the reference data about the stage of bark beetle infestation was obtained through a combination of field survey and visual interpretation of an orthomosaic. The differences of vegetation indices between infested and healthy trees over four time points were statistically evaluated and classified using the Maximum Likelihood classifier. Achieved results confirm our assumptions that it is possible to use a low-cost UAV-based sensor for detection of various stages of bark beetle infestation across seasons; with increasing time after infection, distinguishing infested trees from healthy ones grows easier. The best performance was achieved by the Greenness Index with overall accuracy of 78%–96% across the time periods. The performance of the indices based on near-infrared band was lower.

Resilience and functionality of European Norway spruce forests are increasingly threatened by mass outbreaks of the bark beetle Ips typographus promoted by heat, wind throw and drought. Here, we review current knowledge on Norway spruce and I. typographus interactions from the perspective of drought-stressed trees, host selection, colonisation behaviour of beetles, with multi-level effects of symbiotic ophiostomatoid fungi. By including chemo-ecological, molecular and behavioural perspectives, we provide a comprehensive picture on this complex, multitrophic system in the light of climate change. Trees invest carbon into specialised metabolism to produce defence compounds against biotic invaders; processes that are strongly affected by physiological stress such as drought. Spruce bark contains numerous terpenoid and phenolic substances, which are important for bark beetle aggregation and attack success. Abiotic stressors such as increased temperatures and drought affect composition, amounts and emission rates of volatile compounds. Thus, drought events may influence olfactory responses of I. typographus , and further the pheromone communication enabling mass attack. In addition, I. typographus is associated with numerous ophiostomatoid fungal symbionts with multiple effects on beetle life history. Symbiotic fungi degrade spruce toxins, help to exhaust tree defences, produce beetle semiochemicals, and possibly provide nutrition. As the various fungal associates have different temperature optima, they can influence the performance of I. typographus differently under changing environmental conditions. Finally, we discuss why effects of drought on tree-killing by bark beetles are still poorly understood and provide an outlook on future research on this eruptive species using both, field and laboratory experiments.

In this article, we investigated the detection of forest vegetation changes during the period of 2017 to 2019 in the Low Tatras National Park (Slovakia) and the Sumava National Park (Czechia) using Sentinel-2 data. The evaluation was based on a time-series analysis using selected vegetation indices. The case studies represented five different areas according to the type of the forest vegetation degradation (one with bark beetle calamity, two areas with forest recovery mode after a bark beetle calamity, and two areas without significant disturbances). The values of the trajectories of the vegetation indices (normalized difference vegetation index (NDVI) and normalized difference moisture index (NDMI)) and the orthogonal indices (tasseled cap greenness (TCG) and tasseled cap wetness (TCW)) were analyzed and validated by in situ data and aerial photographs. The results confirm the abilities of the NDVI, the NDMI and the TCW to distinguish disturbed and undisturbed areas. The NDMI vegetation index was particularly useful for the detection of the disturbed forest and forest recovery after bark beetle outbreaks and provided relevant information regarding the health of the forest (the individual stages of the disturbances and recovery mode). On the contrary, the TCG index demonstrated only limited abilities. The TCG could distinguish healthy forest and the gray-attack disturbance phase; however, it was difficult to use this index for detecting different recovery phases and to distinguish recovery phases from healthy forest. The areas affected by the disturbances had lower values of NDVI and NDMI indices (NDVI quartile range Q2–Q3: 0.63–0.71; NDMI Q2–Q3: 0.10–0.19) and the TCW index had negative values (Q2–Q3: −0.06–−0.05)). The analysis was performed with a cloud-based tool—Sentinel Hub. Cloud-based technologies have brought a new dimension in the processing and analysis of satellite data and allowed satellite data to be brought to end-users in the forestry sector. The Copernicus program and its data from Sentinel missions have evoked new opportunities in the application of satellite data. The usage of Sentinel-2 data in the research of long-term forest vegetation changes has a high relevance and perspective due to the free availability, distribution, and well-designed spectral, temporal, and spatial resolution of the Sentinel-2 data for monitoring forest ecosystems.

Research Highlights: Bark beetles are important agents of disturbance regimes in temperate forests, and specifically in a connected wind-bark beetle disturbance system. Large-scale windthrows trigger population growth of the European spruce bark beetle (Ips typographus L.) from endemic to epidemic levels, thereby allowing the killing of Norway spruce trees over several consecutive years. Background and Objectives: There is a lack of evidence to differentiate how outbreaks are promoted by the effects of environmental variables versus beetle preferences of trees from endemic to outbreak. However, little is known about how individual downed-tree characteristics and local conditions such as tree orientation and solar radiation affect beetle colonization of downed trees. Materials and Methods: To answer this question, we investigated the infestation rates and determined tree death categories (uprooted, broken, and stump) in wind-damaged areas in Western Tatra Mts. in Carpathians (Slovakia) from 2014–2016, following a windthrow in May 2014. In total, we investigated 225 trees over eight transects. For every tree, we measured its morphological (tree height, crown characteristics), environmental (solar radiation, terrain conditions, trunk zenith), temporal (time since wind damage), and beetle infestation (presence, location of attack, bark desiccation) parameters. We applied Generalized Additive Mixed Models (GAMM) to unravel the main drivers of I. typographus infestations. Results: Over the first year, beetles preferred to attack broken trees and sun-exposed trunk sides over uprooted trees; the infestation on shaded sides started in the second year along with the infestation of uprooted trees with lower desiccation rates. We found that time since wind damage, stem length, and incident solar radiation increased the probability of beetle infestation, although both solar radiation and trunk zenith exhibited nonlinear variability. Our novel variable trunk zenith appeared to be an important predictor of bark beetle infestation probability. We conclude that trunk zenith as a simple measure defining the position of downed trees over the terrain can anticipate beetle infestation. Conclusions: Our findings contribute to understanding of the bark beetle’s preferences to colonize windthrown trees in the initial years after the primary wind damage. Further, our findings can help to identify trees that are most susceptible to beetle infestation and to prioritize management actions to control beetle population while maintaining biodiversity.

Insect outbreaks affect forests, causing the deaths of trees and high economic loss. In this study, we explored the detection of European spruce bark beetle (Ips typographus, L.) outbreaks at the individual tree crown level using multispectral satellite images. Moreover, we explored the possibility of tracking the progression of the outbreak over time using multitemporal data. Sentinel-2 data acquired during the summer of 2020 over a bark beetle–infested area in the Italian Alps were used for the mapping and tracking over time, while airborne lidar data were used to automatically detect the individual tree crowns and to classify tree species. Mapping and tracking of the outbreak were carried out using a support vector machine classifier with input vegetation indices extracted from the multispectral data. The results showed that it was possible to detect two stages of the outbreak (i.e., early, and late) with an overall accuracy of 83.4%. Moreover, we showed how it is technically possible to track the evolution of the outbreak in an almost bi-weekly period at the level of the individual tree crowns. The outcomes of this paper are useful from both a management and ecological perspective: it allows forest managers to map a bark beetle outbreak at different stages with a high spatial accuracy, and the maps describing the evolution of the outbreak could be used in further studies related to the behavior of bark beetles.

In temperate forests, elevated frequency of drought related disturbances will likely increase the incidence of interactions between disturbances such as bark beetle epidemics and wildfires. Our understanding of the influence of recent drought and insect-induced tree mortality on wildfire severity has largely lacked information from forests adapted to frequent fire. A recent unprecedented tree mortality event in California’s Sierra Nevada provides an opportunity to examine this disturbance interaction in historically frequent-fire forests. Using field data collected within areas of recent tree mortality that subsequently burned in wildfire, we examined whether and under what conditions wildfire severity relates to severity of prefire tree mortality in Sierra Nevada mixed-conifer forests. We collected data on 180 plots within the 2015 Rough Fire and 2016 Cedar Fire footprints (California, USA). Our analyses identified prefire tree mortality as influential on all measures of wildfire severity (basal area killed by fire, RdNBR, and canopy torch) on the Cedar Fire, although it was less influential than fire weather (relative humidity). Prefire tree mortality was influential on two of three fire-severity measures on the Rough Fire, and was the most important predictor of basal area killed by fire; topographic position was influential on two metrics. On the Cedar Fire, the influence of prefire mortality on basal area killed by fire was greater under milder weather conditions. All measures of fire severity increased as prefire mortality increased up to prefire mortality levels of approximately 30–40%; further increases did not result in greater fire severity. The interacting disturbances shifted a pine-dominated system (Rough Fire) to a cedar–pine–fir system, while the pre-disturbance fir–cedar system (Cedar Fire) saw its dominant species unchanged. Managers of historically frequent-fire forests will benefit from utilizing this information when prioritizing fuels reduction treatments in areas of recent tree mortality, as it is the first empirical study to document a relationship between prefire mortality and subsequent wildfire severity in these systems. This study contributes to a growing body of evidence that the influence of prefire tree mortality on wildfire severity in temperate coniferous forests may depend on other conditions capable of driving extreme wildfire behavior, such as weather.

Salvage logging is one of most frequently applied management responses to forest disturbances world‐wide. As forest disturbances are increasing, so too is the application of salvage logging, yet its effects on ecosystems remains incompletely understood. In the Norway spruce (Picea abies (L.) Karst.) forests of Europe, salvaging of windfelled trees is inter alia applied to reduce the risk of bark beetle outbreaks (mainly Ips typographus L.). By preventing further disturbances, salvage logging can conserve live tree carbon (C) in forest landscapes. At the same time salvage logging reduces C stocks in detrital pools via the extraction of disturbed trees, its net effect thus remains unclear. We used the forest landscape model iLand to explore the effect of a wide range of salvaging intensities on (a) subsequent bark beetle outbreaks, and (b) landscape‐scale forest C stocks in a Norway spruce‐dominated production forest in Slovakia under past and future climatic conditions. Climate change resulted in a two‐ to three‐fold increase in bark beetle disturbances throughout the 21st century in our simulations. We found that removing >95% of disturbed trees can effectively buffer the effect of increasing disturbances, dampening bark beetle infestations and increasing live tree C. Total ecosystem C followed a U‐shaped pattern over salvaging intensity, with highest values in no salvage and 100% salvage scenarios. However, realistic rates of salvaging (<95% of disturbed trees detected and removed) had no significant effect on bark beetle dynamics and live tree C, and reduced the total ecosystem C stored in the landscape. Furthermore, the effect of reduced bark beetle disturbance under intensive salvaging was partly offset by increased wind disturbance. Synthesis and applications. Clearing disturbed areas to prevent future disturbances from bark beetles and conserve live tree carbon should only be applied where very high salvaging rates are feasible (i.e. small and concentrated disturbances). Considering that changing disturbance regimes make high‐intensity salvaging increasingly challenging, alternative disturbance management approaches need to be developed. Foreign Language Czech Odstraňování napadených či poškozených stromů (tzv. nahodilé těžby) je nejrozšířenější reakcí na výskyt různých disturbancí v lesních ekosystémech. Klimatická změna vede k intenzívnějšímu vlivu disturbancí na lesy a je tedy zapotřebí očekávat i nárůst rozsahu a intenzity nahodilých těžeb. Ve smrkových porostech v Evropě se nahodilé (v tomto případě přesněji sanitární) těžby větrem poškozených stromů provádějí proto, aby se zabránilo náletu lýkožroutů na vhodný rozmnožovací materiál či snížil nárůst populace, která již polámané stromy obsadila. Pokud zabráníme přemnožením kůrovců, uchováme rovněž zásobu uhlíku v živých stromech. Na druhé straně, těžbami odstraňujeme část uhlíku uloženého v mrtvém dřevu. Výsledný efekt těchto dvou protichůdných procesů na celkovou uhlíkovou bilanci však není znám. V této studii jsme pomocí ekosystémového modelu iLand hodnotili vliv různých intenzit odstraňování větrem poškozených stromů na (a) následné přemnožení kůrovce a (b) na celkovou zásobu uhlíku v ekosystému. Výzkum byl realizován v hospodářském lese s dominancí smrku (75%) s rozlohou 16 050 ha. V důsledku změny klimatu došlo v průběhu 21. století k dvou až trojnásobnému nárůstů objemu stromů napadených kůrovcem. Odstranění více než 95% větrem poškozených stromů účinně zmírnilo intenzitu kůrovcových přemnožení a došlo tím ke zvýšení zásob uhlíku v živých stromech. Závislost celkové zásoby uhlíku v ekosystémů na různé intenzitě těžeb měla tvar U. Nejvyšší úrovně byly dosahovány při absenci těžeb (uhlík se více kumuluje v mrtvém dřevu) a při celkovém odstranění větrem poškozených stromů (kůrovec je potlačen a uhlík se kumuluje v živých stromech). Realistická intenzita sanitárních těžeb (detekce a odstranění <95% poškozených stromů) neměla na dynamiku kůrovce a zásobu uhlíku v živých stromech téměř žádný vliv, naopak způsobila pokles celkové zásoby uhlíku v ekosystému. Intenzivní prevence vzniku přemnožení kůrovce zároveň vyvolala v dlouhodobém horizontu nárůst poškození porostů větrem a částečně tak negovala snahu o prevenci disturbancí. Z těchto výsledků vyplývá, že odstraňování větrem poškozených stromů s cílem prevence nebo potlačení přemnožení kůrovce a uchování zásob uhlíku má smysl realizovat pouze tehdy, pokud je možné dosáhnout velice vysokou intenzitu těžeb (např. při menších a soustředěných polomech). Protože s klimatickou změnou dochází k zesílení vlivu disturbancí a změně dlouhodobých disturbančních režimů, provádění vysokých intenzit sanitárních těžeb bude stále obtížnější. Je proto zapotřebí hledat alternativní způsoby managementu hospodářských lesů orientující se na podporu jejich přirozené rezilience. Clearing disturbed areas to prevent future disturbances from bark beetles and conserve live tree carbon should only be applied where very high salvaging rates are feasible (i.e. small and concentrated disturbances). Considering that changing disturbance regimes make high‐intensity salvaging increasingly challenging, alternative disturbance management approaches need to be developed.