Estimating the most effective and economical pheromone for monitoring the European spruce bark beetle

In recent decades, there have been an increasing number of outbreaks of Ips typographus in Europe. A large amount of sanitary felling has taken place, with significant economic and ecological consequences. In order to anticipate such large-scale outbreaks, an effective monitoring system should be set up. One important aspect of monitoring is deciding which pheromone to use. Therefore, we decided to test five different commercially available pheromone lures under different disturbance conditions: Pheroprax®, IT Ecolure Extra®, Ipstyp®, Ipsowit® and Typosan®. We investigated the ability of the pheromones to distinguish between disturbed and undisturbed locations, their cost-efficiency ratio, and side effects such as bycatch abundance and composition. We set 50 traps in two areas with sites that were disturbed and undisturbed by windstorms. We collected the catch from traps every 1–2 weeks from the end of March until the end of September in 2019. We found that IT Ecolure Extra®, Ipsowit® and ® Pheroprax® caught the most I. typographus and best showed changes in the trap catch of I. typographus throughout the whole season. There was a low amount of bycatch (<6% of the total catch) and a low number of predators (a few specimens), but some groups seem to prefer certain pheromones. The cost of the pheromones increased with their effectiveness. However, pheromone costs are low relative to the personnel costs involved in setting traps and collecting the catch. Based on all of the gathered data, we created an index which helps to assess the cost-efficiency of the five chosen commercially available pheromones. We also present guidelines on how to make such an index to assist other researchers in choosing the right pheromone for monitoring populations of I. typographus or other bark beetle species.     

Characterization of microsatellite loci in the spruce bark beetle Ips typographus (Coleoptera: Scolytinae)

Ips typographus is an economically important pest of Norway spruce stands. We developed five polymorphic microsatellite markers using a biotin enrichment protocol. The number of alleles ranged from three to 11 per locus and no strong evidence for null alleles was found. Heterozygosity ranged from 0.3 to 0.97. These markers could be useful tools to study the population structure and genetic consequences of I. typographus population outbreaks.     

Colonization of storm gaps by the spruce bark beetle: influence of gap and landscape characteristics

• 1 After storm disturbances, there is a risk for degradation of the quality of fallen trees, and for subsequent tree mortality caused by the spruce bark beetle Ips typographus (L.) (Coleoptera: Curculionidae). Models assessing the risk for bark beetle colonization of different kinds of storm gaps would be a valuable tool for management decisions.
• 2 The present study aimed to determine which gap and landscape characteristics are correlated with the probability of colonization of wind‐felled Norway spruce trees by I. typographus.
• 3 The study included 36 storm gaps, varying in size from three to 1168 wind‐felled spruces, created by the storm Gudrun in southern Sweden in January 2005.
• 4 In the first summer, on average, 5% of the wind‐felled spruces were colonized by I. typographus. The percentage of colonized wind‐felled trees per gap was negatively correlated with the total area of storm gaps within 2000 m in the surrounding forest landscape.
• 5 In the second summer, the proportion of colonized trees increased to 50%. Both gap (mean diameter of wind‐felled trees and basal area of living spruce trees) and landscape variables (amount of spruce forest) were significantly correlated with colonization percentage and explained almost 50% of the variation between gaps.
• 6 There was no relationship between gap area and colonization percentage. This implies that landscapes with many large storm gaps, where logging resources will be most effectively used, should be salvaged first.

     

Escape in space from enemies: a comparison between stands with and without enhanced densities of the spruce bark beetle

1 Populations of the spruce bark beetle, Ips typographus (L.), are known to grow rapidly in storm‐disturbed stands as a result of relaxation from intraspecific competition. In the present study, it was tested whether a second mechanism, escape in space from natural enemies, also contributes to the rapid population increase. 2 The experiment was conducted during the initiation phase of five local outbreaks of I. typographus triggered by a storm‐disturbance in November 1995 in southern Sweden. 3 The impact of natural enemies on the ratio of increase (number of daughters per mother) of I. typographus was compared pairwise between disturbed stands with high numbers of storm‐felled trees and undisturbed stands without wind‐felled trees. 4 Enemy impact was assessed by comparing the ratio of increase in uncaged (exposed to enemies) and caged (protected from enemies) bolts colonized by I. typographus prior to being placed in the stands. The experiment was conducted in the second and third summers after the storm‐felling. 5 Enemy impact was about twice as high in stands without wind‐felled trees compared with in stands with wind‐felled trees in the second summer whereas there was no significant difference between the stand types in the third summer. 6 The result demonstrates that spatial escape from enemies contributes to the rapid population growth of I. typographus after storm‐disturbances.     

Tree killing by Ips typographus (Coleoptera: Scolytidae) at stand edges with and without colonized felled spruce trees

Abstract 1 To maintain biodiversity in managed spruce forests in Sweden more wind‐felled trees must be retained. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col. Scolytidae). 2 To simulate wind‐felled trees, living spruce trees were cut at spruce stand edges bordering fresh clear‐cuttings. Treatments comprised edges with zero, one or five cut trees colonized by I. typographus. Edges with naturally wind‐felled trees colonized by I. typographus were also included in the analyses. 3 During the two following summers, the number of trees killed by I. typographus did not differ between edges with and without felled trees, or between edges with one or five felled trees. 4 Within edges with felled trees, there were more killed trees close to the felled trees than at other parts of the edges. Thus, felled trees provided focal points for attacks within edges. 5 It is concluded that small numbers of wind‐felled trees colonized by I. typographus may be left near spruce stand edges without increasing the risk of beetle‐induced tree mortality.     

Gallery construction and oviposition of the bark beetle Ips typographus (Coleoptera: Scolytidae) at different breeding densities

Abstract. 1. The bole of a Norway spruce, Picea abies (L.) Karst., was cut in 25 cm sections and infested in the laboratory with 2 or 8 females/dm² of Ips typographus (L.) and an equal number of males.
2. One log of each beetle density was dissected every second day, beginning 2 days after female introduction.
3. The oviposition rate was highest at the beginning of the breeding period for both densities and then declined, especially at the high density. Residence time is a useful approximation for the number of laid eggs in the beginning of the oviposition period or when the density is low.
4. Final gallery length and egg number per gallery were on average significantly smaller at density 8 than at density 2.
5. Linear relationships between total gallery length and number of egg niches were found to be similar for the two densities.
6. An estimate of the egg to adult mortality at different densities was found by using data from a field study that related gallery length to breeding density in attacked living trees, together with published relationships between density and egg niches or emergence of new adults.
7. It was concluded that both decreased oviposition rate and earlier re-emergence at higher densities contribute to a declining oviposition with increasing density. The final offspring production is futher reduced by lower survival due to larval competition as density increases.     

A predictive framework to assess spatio‐temporal variability of infestations by the European spruce bark beetle

Natural disturbances are key factors for the development of forest ecosystems. In forests of central Europe and Scandinavia, the European spruce bark beetle Ips typographus is the most devastating biotic disturbance agent in Norway spruce Picea abies, but our understanding of the factors determining its spatio‐temporal dynamics is still quite limited. To quantify the drivers of bark beetle dynamics, we analyzed a survey dataset with annual resolution that covers 17 yr and 469 forest districts (10 860 km² of forest area) all over Switzerland. We used Poisson log‐normal models in a Bayesian framework to analyze the spatio‐temporal dynamics of bark beetle infestation spots at the forest district level. Bark beetle infestations increased with increasing heat sum (> 8.3°C), volume of standing Norway spruce stock, and the number of infestation spots of the previous year. Precipitation tended to slightly affect the risk of bark beetle infestations. Two major storm events further increased the spatio‐temporal variability of bark beetle infestations. Spruce abundance, storm damage and temperature are known to be important factors influencing the population dynamics of the European spruce bark beetle. Our study is the first to quantify the combined effects of spruce abundance and heat sum, whereby the heat sum turned out to be the most important and consistent predictor. Because our study area encompasses large ecological and climatological gradients, our model is likely to be applicable to Norway spruce forests in other regions of central Europe and Scandinavia.     

Attacks on living spruce trees by the bark beetle Ips typographus (Col. Scolytidae) following a storm-felling: a comparison between stands with and without removal of wind-felled trees

Abstract 1 To maintain biodiversity in forests more wind‐felled trees must be left where they fall. However, there is concern among forest owners that this may result in higher tree mortality caused by the spruce bark beetle, Ips typographus (L.) (Col.: Scolytidae). 2 In the 5 years following a major storm disturbance the number of standing spruces killed by I. typographus was determined in a total of 53 stands. In five of the stands all wind‐thrown trees were left (unmanaged stands) and in 48 of the stands, which were situated at distances of 1.4–10.0 km from each focal unmanaged stand, the wind‐felled trees were removed directly after the storm (managed stands). In the winter preceding the fifth summer new storm‐fellings occurred in the study area. 3 In the 4‐year period between the first and second storm‐fellings, 50–322 standing trees were killed by I. typographus per unmanaged stand. There was a direct linear relationship between the number of storm‐felled spruces colonized by I. typographus and the number of trees subsequently killed in the unmanaged stands. 4 Tree mortality caused by I. typographus in the unmanaged stands was almost nil in the first year, peaked in the second or third year, and decreased markedly to a low level in the fourth and fifth years. 5 In the 4‐year period between the first and second storm‐fellings twice as many trees were killed per ha in the unmanaged stands than in the managed stands: the average difference being 6.2 killed trees per ha, equivalent to 19% of the number of spruce trees felled by the first storm in the unmanaged stands. 6 Much higher numbers of trees were killed per ha in the stand edges than in the interiors of both the unmanaged and the managed stands.     

Prolonged development time of the bark beetle predator Thanasimus formicarius (Col.: Cleridae) in relation to its prey species Tomicus piniperda (L.) and Ips typographus (L.) (Col.: Scolytidae)

1 The generation time of the bark beetle predator Thanasimus formicarius (L.) (Col.: Cleridae) was found to be predominantly two years both in the field and in rearing experiments conducted with two of its main prey species, the pine shoot beetle Tomicus piniperda (L.) and the spruce bark beetle Ips typographus (L.) (Col.: Scolytidae). 2 Emergence of T. formicarius adults in the first summer was only observed in one of the two rearing experiments, and these individuals represented only 6% of that generation. 3 All individuals not emerging as adults in the first summer remained as larvae in their pupal chambers until the second summer. Pupae were found starting around mid-June, and adults (in pupal chambers) were found from late July through to the end of August. 4 Newly emerged adults had to feed in order to survive hibernation. 5 The existence of T. formicarius races, specialized on certain bark beetle species and with phenologies matching their hosts, could not be demonstrated. After hibernation there was no difference in feeding activity, timing of egg-laying or proportion of egg-laying females between the T. formicarius adults reared as larvae on T. piniperda (flight period in April) and those reared as larvae on I. typographus (main flight period generally starting in late May or early June) when exposed to a temperature and day-length typical of the early spring conditions prevailing during the flight period of T. piniperda. 6 T. formicarius was parasitized by Enclisis vindex (Tschek) (Hym.: Ichneumonidae) in the pupal chamber. 7 The importance of these findings for the population dynamics of bark beetles is discussed.     

Field response of the northern spruce bark beetle Ips duplicatus (Sahlberg) (Coleoptera: Curculionidae,Scolytinae) to different combinations of synthetic pheromone with (−)‐α‐pinene and (+)‐limonene

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Temperature‐dependent development of the great European spruce bark beetle Dendroctonus micans (Kug.) (Coleoptera: Curculionidae: Scolytinae) and its predator Rhizophagus grandis Gyll. (Coleoptera: Monotomidae: Rhizophaginae)

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Water-macromolecule interactions by NMR: A quadrature-free constant-time approach and its application to CI2

A pulse sequence is proposed to select water magnetization with enhanced specificity through a synergetic combination of several filtering principles. This approach relies on a constant-time evolution period implemented without quadrature detection, which results in a 2 increase in signal-to-noise ratios as compared to traditional non-selective methods for water filtration. In addition, the quadrature-free constant-time block facilitates the implementation of the water flip-back strategy, which leads to further gains in sensitivity. The proposed experiment was applied to unlabeled HEW lysozyme and to ¹⁵ N-labeled chymotrypsin inhibitor 2 which was partially or non ¹³C-enriched. Water molecules belonging to a spine of hydration between two pseudo -sheet strands were identified, solving previously reported discrepancies between the X-ray and the refined NMR structure of CI2. The proposed experiment is particularly suitable for hydration studies of mixtures of labeled and unlabeled components, such as ligand–macromolecule complexes.     

Flood mapping with remote sensing and hydrochemistry: A new method to distinguish the origin of flood water during floods

River flooding is important for the ecological functioning of river floodplains. It is implicitly assumed that in many river floodplains during floods, river water is spreading all over the floodplain. We hypothesize that during flood events a spatial distribution of water types exists, which is correlated to different water sources (river water, atmospheric water and groundwater) and to the spatial distribution of vegetation types. The objective of this paper is to assess a new methodology to determine the extent of flooding and the spatial distribution of different water sources during the flood, using GPS, multispectral remote sensing and hydrochemical analyses. This methodology is applied to the Biebrza River Lower Basin, which has little human impact. Remote sensing resulted in a map distinguishing inundated areas from dry areas, which showed 85% agreement with GPS field measurements. Principal Component Analyses and Cluster Analyses on the measured water chemistry identified different water sources during the flood (river water, groundwater, rainwater) and showed the effects of human impact on water quality. River flood water dominated the entire inundation zone in the northern Lower Basin, which is narrower and steeper than the southern Lower Basin where groundwater and rainwater were significant contributors to the major part of the inundated area. Vegetation in the river flood zone is distinctly different from the rest of the floodplain. Due to mixing of ground- and rainwater, correlation analyses between vegetation and water type were not possible outside the river flood zone. The new methodology is effective in distinguishing inundated areas from dry regions and in separating river flood water from other water sources during a flood.     

A modelling approach to estimate the effect of exotic pollinators on exotic weed population dynamics: bumblebees and broom in Australia

The role of mutualisms in contributing to species invasions is rarely considered, inhibiting effective risk analysis and management options. Potential ecological consequences of invasion of non‐native pollinators include increased pollination and seed set of invasive plants, with subsequent impacts on population growth rates and rates of spread. We outline a quantitative approach for evaluating the impact of a proposed introduction of an invasive pollinator on existing weed population dynamics and demonstrate the use of this approach on a relatively data‐rich case study: the impacts on Cytisus scoparius (Scotch broom) from proposed introduction of Bombus terrestris. Three models have been used to assess population growth (matrix model), spread speed (integrodifference equation), and equilibrium occupancy (lattice model) for C. scoparius. We use available demographic data for an Australian population to parameterize two of these models. Increased seed set due to more efficient pollination resulted in a higher population growth rate in the density‐independent matrix model, whereas simulations of enhanced pollination scenarios had a negligible effect on equilibrium weed occupancy in the lattice model. This is attributed to strong microsite limitation of recruitment in invasive C. scoparius populations observed in Australia and incorporated in the lattice model. A lack of information regarding secondary ant dispersal of C. scoparius prevents us from parameterizing the integrodifference equation model for Australia, but studies of invasive populations in California suggest that spread speed will also increase with higher seed set. For microsite‐limited C. scoparius populations, increased seed set has minimal effects on equilibrium site occupancy. However, for density‐independent rapidly invading populations, increased seed set is likely to lead to higher growth rates and spread speeds. The impacts of introduced pollinators on native flora and fauna and the potential for promoting range expansion in pollinator‐limited ‘sleeper weeds’ also remain substantial risks.     

A probabilistic approach to the use of pollen indicators for plant attributes and biomes: an application to European vegetation at 0 and 6 ka

Aim This paper presents a probabilistic method for the characterization of pollen taxa using attributes, and for the reconstitution of past biomes. The probabilities are calculated on the basis of European floristic and pollen databases sufficiently large and exhaustive to provide robust estimates. Location The analysis is based on data from approximately 1000 sites throughout Europe. Method We use all the pollen data from the European Pollen Database (EPD), which contains about 50 000 pollen assemblages distributed across Europe and covering the period from the Last Glacial Maximum to the present. Using existing floras, each pollen taxon has been characterized by allocating one or more modes of several attributes, chosen according to the biogeography and phenology of the taxon. With this information, conditional probabilities are defined, representing the chance of a given attribute mode occurring in a given pollen spectrum, when the taxa assemblage is known. The concept of co‐occurrence is used to provide a greater amount of information to compensate for difficulties in the identification of pollen grains, allowing a better interpretation when there is little diversity in the pollen assemblage. Results The method has been validated using a dataset of modern samples against existing methods of biome classification and remote sensing data. An application is proposed in which the new method is used to produce biomes for pollen data 6000 years ago. This confirms previous results showing an extension of the deciduous forest to the north, east and south, explained by milder winters in western and northern Europe, and cooler and wetter climate in the Mediterranean region. Conclusion The results show the new method to be efficient, reliable and flexible and to be an improvement over the previous method of biomization. They will be used to test simulations of earth system models running on periods with climate significantly different from the present day, enabling a robust test of the validity of applying these models to the future.     

Position of residues in transmembrane peptides with respect to the lipid bilayer: A combined lipid NOEs and water chemical exchange approach in phospholipid bicelles

The model transmembrane peptide P16 (Ac-KKGLLLALLLLALLLALLLKKA-NH₂) was incorporated into small unaligned phospholipid bicelles, which provide a `native-like' lipid bilayer compatible with high-resolution solution NMR techniques. Using amide-water chemical exchange and amide-lipid cross-relaxation measurements, the interactions between P16 and bicelles were investigated. Distinctive intermolecular NOE patterns observed in band-selective 2D-NOESY spectra of bicellar solutions with several lipid deuteration schemes indicated that P16 is preferentially interacting with the `bilayered' region of the bicelle rather than with the rim. Furthermore, when amide-lipid NOEs were combined with amide-water chemical exchange cross-peaks of selectively ¹⁵N-labeled P16 peptides, valuable information was obtained about the position of selected residues relative to the membrane-water interface. Specifically, three main classes were identified. Class I residues lie outside the bilayer and show amide-water exchange cross-peaks but no amide-lipid NOEs. Class II residues reside in the bilayer-water interface and show both amide-water exchange cross-peaks and amide-lipid NOEs. Class III residues are embedded within the hydrophobic core of the membrane and show no amide-water exchange cross-peaks but strong amide-lipid NOEs.     

A 3-Mbp fragment on rat chromosome 1 affects susceptibility both to stroke and kidney injury under salt loading in the stroke-prone spontaneously hypertensive rat: a genetic approach using multiple congenic strains

We have previously reported that a major quantitative trait locus (QTL) responsible for susceptibility to salt-induced stroke in the stroke-prone spontaneously hypertensive rat (SHRSP) is located in a 3-Mbp region on chromosome 1 covered by SHRSP.SHR-(D1Rat23-D1Rat213)/Izm (termed Pr1.31), a congenic strain with segments from SHRSP/Izm introduced into the stroke-resistant SHR/Izm. Here, we attempted to narrow down the candidate region on chromosome 1 further through analyses of subcongenic strains constructed for the target region. Simultaneously, salt-induced kidney injury was evaluated through the measurement of urinary albumin and the gene expression of renal tubular injury markers (Kim-1 and Clu) to explore a possible mechanism leading to the onset of stroke. All subcongenic strains examined in this study showed lower susceptibility to salt-induced stroke than SHRSP. Interestingly, Pr1.31 had the lowest stroke susceptibility when compared with newly constructed subcongenic strains harboring fragments of the congenic sequence in Pr1.31. Although Kim-1 and Clu expression after 1 week of salt loading in Pr1.31 did not differ significantly from those in SHRSP, the urinary albumin level of Pr1.31 was significantly lower than those of the other subcongenic strains and that of SHRSP. The present results indicated that, although the congenic fragment in Pr1.31 harbored the gene(s) related to salt-induced organ damages, further genetic dissection of the candidate region was difficult due to multiple QTLs suggested in this region. Further analysis using Pr1.31 will unveil genetic and pathophysiological mechanisms underlying salt-induced end organ damages in SHRSP.