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.     

The bark beetle Pityogenes chalcographus (L.) (Scolytidae) in living trees: reproductive success, tree mortality and interaction with Ips typographus

Abstract:  Tree-killing ability of Pityogenes chalcographus was experimentally investigated by baiting living spruce Picea abies with synthetic aggregation pheromone, thereby exposing them to beetle attack. Reproductive success was estimated by rearing beetles from stem sections taken from killed trees. The presence of Ips typographus on several killed trees allowed a comparison of reproductive success in the presence and absence of this species. Furthermore, the possibility that breeding material colonized by P. chalcographus may induce subsequent attacks by I. typographus on nearby trees was also studied. The tree killing ability of P. chalcographus was low as only 8% of the baited trees, attacked by P. chalcographus alone, were killed. Tree death was strongly related to the presence of I. typographus , which is in line with the association with aggressive wood-living fungi in this species. The reproductive success was low, with on average less than one daughter beetle produced per mother beetle. The success was significantly higher in trees also inhabited by I. typographus . Tree-parts colonized by P. chalcographus induced almost no attacks by I. typographus on nearby trees.     

Effect of different semiochemicals blends on spruce bark beetle,Ips typographus (Coleoptera: Scolytidae)

The spruce bark beetle, Ips typographus, is a recent new introduction to the Qilian Mountains of China. An outbreak of these beetles has infested over 0.03 million hectares of spruce forests in this area. Although primary attraction to volatiles has been clearly demonstrated for I. typographus, the existence and role of attraction to insect‐produced pheromones have been widely debated. Currently, commercial lures for I. typographus include only the volatiles ipsdienol, cis‐verbenol, trans‐verbenol, 2‐methyl‐3‐buten‐2‐ol and 2‐phenylethanol in Europe. Several potential pheromone candidates have been identified for I. typographus. Our GC–MS and GC–FID analyses volatiles from hindgut extracts of I. typographus in different attack phases demonstrated that the 2‐methyl‐3‐buten‐2‐ol, ipsdienol, cis‐verbenol and trans‐verbenol as major hindgut components, and ipsenol, 2‐phenylethanol, trans‐ myrtenol and verbenone as minor components. We tested various combinations of semiochemical candidates, to determine an optimal blend. Our results suggest that addition of 2‐methyl‐3‐buten‐2‐ol to either ipsenol alone, or to blends of ipsenol and other semiochemical candidates, significantly enhanced attraction of I. typographus. Therefore, a simple lure consisting of ipsenol and 2‐methyl‐3‐buten‐2‐ol would be an optimal blend of I. typographus in the Qilian Mountains, China. We conclude that this optimal semiochemical blend may provide an effective biological pest control method for use in forest ecosystem against I. typographus.     

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.     

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.     

Occurrence of Ips typographus (Col., Scolytidae) along an urbanization gradient in Brussels, Belgium

Abstract 1 The distribution and dynamics of insect populations in cities is poorly understood. One approach to address this question is to explore the permeability of the urban habitat to species from surrounding rural areas, which can serve as reservoirs in source‐sink dynamics. 2 Here, we present data on the distribution of the forest insect pest of spruce, Ips typographus (Coleoptera, Scolytidae), along two axes entering the city of Brussels (Belgium) from the south‐east and to the town centre. 3 The insect was caught everywhere along these transects, even in heavily urbanized surroundings, and sometimes in relatively high numbers. The catches were highest near the middle of the transects and lower at both ends of them. 4 This pattern was associated, on the one hand, with an urbanization gradient with the numbers of flying individuals increasing with the distance from the city centre and, on the other hand, with lower catches at the periphery of the city where a high proportion of broadleaved trees may have disrupted the response to aggregation pheromones. 5 In addition to the probable rural origin of the beetles, high catches at the Port of Brussels indicated that some of the insects might be of foreign origin and enter the city with imported timber, highlighting a pathway for unintentional introductions of organisms, including potentially invasive species.     

Performance of the tree‐killing bark beetles Ips typographus and Pityogenes chalcographus in non‐indigenous lodgepole pine and their historical host Norway spruce

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Phylogeography and postglacial colonization routes of Ips typographus L. (Coleoptera, Scolytidae)

Ips typographus populations were analysed by enzyme electrophoresis and by sequence analysis in order to quantify the degree of population differentiation. Enzyme electrophoresis showed a high gene flow among all European populations. Analysis of single loci showed that aspartate aminotransferase-2 ( Aat-2 ) clustered the Scandinavian populations apart from the other populations whereas other enzyme loci showed no significant pattern. Analysis of mitochondrial DNA revealed eight haplotypes. The populations from Italy, Croatia and Belgium were the most polymorphic. Two haplotypes (I and II) were found in the majority of Central European populations. In the northern populations only one haplotype (I) was found. The Russian and Lithuanian populations had a particular haplotype (IV) that was not found in any of the other populations. In view of these data it is suggested that the population structure of I. typographus has been influenced by events which took place during and after the last ice age. Populations were forced into refugial areas in the south and in the area north of Moscow along with the host tree, Picea abies . P. abies had four refugial areas: the Apennine Alps, the Dinaric Alps, the Carpathian Alps and Kostroma, an area north of Moscow. With amelioration of the temperature, beetles spread after P. abies . According to the data presented here, I. typographus migrated from the Apennine and/or Dinaric Alps to the north. Whilst there is evidence for high gene flow among populations, bottleneck effects can still be detected in the north and also in the east. Beetles from the refugial area north of Moscow did not spread to Scandinavia or to the west like the host tree.     

Climate change alters elevational phenology patterns of the European spruce bark beetle (Ips typographus)

The European spruce bark beetle Ips typographus is the most important insect pest in Central European forests. Under climate change, its phenology is presumed to be changing and mass infestations becoming more likely. While several studies have investigated climate effects across a latitudinal gradient, it remains an open question how phenology will change depending on elevation and topology. Knowing how an altered climate is likely to affect bark beetle populations, particularly across diverse topographies and elevations, is essential for adaptive management. We developed a time‐varying distributed delay model to predict the phenology of I. typographus. This approach has the particular advantage of capturing the variability within populations and thus representing its stage structure at any time. The model is applied for three regional climate change scenarios, A1B, A2 and RCP3PD, to the diverse topography of Switzerland, covering a large range of elevations, aspects and slopes. We found a strong negative relationship between voltinism and elevation. Under climate change, the model predicts an increasing number of generations over the whole elevational gradient, which will be more pronounced at low elevations. In contrast, the pre‐shift in spring swarming is expected to be greater at higher elevations. In comparison, the general trend of faster beetle development on steep southern slopes is only of minor importance. Overall, the maximum elevation allowing a complete yearly generation will move upwards. Generally, the predicted increase in number of generations, earlier spring swarming, more aggregated swarming, together with a projected increase in drought and storm events, will result in a higher risk of mass infestations. This will increase the pressure on spruce stands particularly in the lowlands and require intensified management efforts. It calls for adapted long‐term silvicultural strategies to mitigate the loss of ecosystem services such as timber production protection against rockfall and avalanches and carbon storage.     

A host monoterpene influences Ips typographus (Coleoptera: Curculionidae,Scolytinae) responses to its aggregation pheromone

1 Host tree terpenes can influence attraction of conifer‐infesting bark beetles to their aggregation pheromones, and both synergistic and inhibitory effects have been reported. 2 We tested a gradient of ratios of (–)‐α‐pinene, the predominant monoterpene in Norway spruce, to the pheromone of Ips typographus, a major pest of Norway spruce. 3 Attraction of I. typographus increased as the release rate of (–)‐α‐pinene increased. The two highest (–)‐α‐pinene : pheromone ratios (526 : 1 and 2595 : 1) attracted twice as many I. typographus as pheromone alone, whereas low to intermediate ratios (56 : 1, 274 : 1) did not differ from pheromone alone. 4 Our results are in agreement with a proposed model, which suggests that bark beetles display unique response profiles to host terpenes depending on the physiological condition of the host trees that they typically colonize. Ips typographus, which is an aggressive species capable of colonizing and killing healthy trees, showed an increased attraction to monoterpene : pheromone ratios, and this may be high enough to inhibit attraction of less aggressive beetle species typically colonizing dead, dying or stressed trees. 5 Attraction of associates of I. typographus was also modified by (–)‐α‐pinene. Ips duplicatus, a competitor of I. typographus, showed increased attraction to the pheromone of I. typographus across all concentrations of (–)‐α‐pinene.     

Size and shape changes of Ips typographus L. (Coleoptera: Scolytinae) in relation to population level

1 High intraspecific competition is known to occur during Ips typographus outbreaks, and is thought to be the main factor regulating epidemic populations by affecting beetle population productivity. However, little is known about the consequences of intraspecific competition on population quality during outbreaks, although it could have consequences on beetle population dynamics. 2 Ips typographus morphological variations among localities, years and beetle population levels were investigated in 10 Norway spruce (Picea abies) stands having various beetle damage intensities. Beetle size and shape estimators based on wing length, and using isometric size and log‐shape ratios, were employed. Field‐caught beetles were compared with beetles emerging from controlled breeding at different densities, performed in the laboratory. Beetles from this colony were also used to check the influence of breeding densities on the size estimator. 3 Size variations occurred among localities and years and were consistent with the epidemic or latent status of the beetle populations. Controlled breeding confirmed the negative effect of beetle densities encountered in the field on offspring size. Two hypotheses are formulated to explain this increase of intraspecific competition during an outbreak, but our data support the effect of host quality change between latent and epidemic populations. 4 Shape variations also occurred among localities but were unrelated to beetle population levels. No groups consistent with a geographical structure were found, suggesting low genetic variation for I. typographus populations in France.     

Recapture of Ips typographus L. (Col., Scolytidae) with attractants of low release rates: localized dispersion and environmental influences

1 The dispersal of Ips typographus L. (Col., Scolytidae) was studied using a mark–release–recapture approach in a grid of traps equipped with pheromone lures of release rates of about 8.4 mg/day of 2‐methyl‐3‐buten‐2‐ol (MB) and 0.29 mg/day of (S)‐cis‐verbenol (cV) in experiment 1, and 1.2 mg/day of MB and 0.04 mg/day of cV in experiment 2. 2 We investigated whether beetle dispersal reflected the simple diffusion pattern observed in previous I. typographus experiments, for which attractant release rates generally approached 50 mg/day of MB and 1 mg/day of cV. We also examined how environmental parameters (wind) and human activities (felling) could influence the beetles' flight. 3 The recapture percentage was higher in experiment 1 than in experiment 2: respectively, 7.0% (with 64 traps) and 2.3% (with 100 traps) of the beetles that took off were caught in the traps. 4 With the higher release rate (experiment 1), trap catches decreased with increased distance, whereas with the lower release rate (experiment 2), trap catches rose between 50 and 100 m then decreased with increasing distance. 5 Flight was little orientated by prevailing wind directions, a feature probably explained by the low wind speeds (0–1.2 m/s) observed throughout the study. 6 High trap catches of unmarked beetles close to areas undergoing thinning activities suggest that the presence of freshly cut spruce and larch material could have an influence on dispersal, attracting the beetles into the felling area. Spatial analyses show that capture patterns were autocorrelated up to distances of about 250 m.     

Metabolism of nitrate and ammonium in seedlings of Norway spruce (Picea abies) measured by in vivo 14N and 15N NMR spectroscopy

In vivo ¹⁵N and ¹⁴N nuclear magnetic resonance spectroscopy was used to investigate the assimilation of nitrate and ammonium in seedlings of Norway spruce (Picea abies [L.] Karst.). The main objective was to study accumulation of free NH+₄ and examine to what extent the nitrogen source affects the composition of the free amino acid pools in roots, stems and needles. NH+₄ concentrations in plants growing in the presence of 0.5–50 mM ammonium were quantified using ¹⁴N NMR. The NH+₄ values in tissues ranged from 6 to 46 μmol (g fresh weight)^?1. with highest concentrations in roots and needles. The tissue NH+₄ peaked at 5.0 mM NH+₄ in the medium. and failed to increase when NH+₄ in the medium was increased to 50 mM, indicating metabolic control of the concentration of this cation in tissues. The ¹⁴N NMR spectra were used to estimate pH of the NH+₄ storage pools. Based on the pH sensitivity of the quintet of ¹⁴NH+₄ resonance, we suggest that the pH of the ammonium storage compartments in the roots and stems should be 3.7–3.8, and in needles 3.4–3.5, representing extremely low pH values of the tissue. ¹⁵N from nitrate or ammonium was first incorporated into the amide group of glutamine and then into α-amino groups, confirming that the glutamine synthetase/ glutamate synthase cycle is the major route of nitrogen assimilation into amino acids and thus plays a role in lowering the levels of NH+₄ in the cytoplasm. NH+₄ can also be assimilated in roots in plants growing in darkness. The main ¹⁵N-labelled amino acids were glutamine. arginine and alanine. Almost no ¹⁵N signals from needles were observed. Double labelling (δN + w, wN) of arginine is consistent with the operation of the ornithine cycle, and enrichment indicates that this cycle is a major sink of newly assimilated nitrogen. Nitrogen assimilation in roots in the presence of added methionine sulphoximine and glutamate indicated the catabolic action of glutamate dehydrogenase. The ¹⁵N NMR spectra of plants grown on ¹⁵N-urea showed a marked increase in the labelling of ammonium and glutamine. indicating high urease activity. Amino acids were also quantified using high pressure liquid chromatography. Arginine was found to be an important transport form of nitrogen in the stem.     

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.     

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.

     

Reducing spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) emergence for hibernation in central British Columbia by felling infested trees

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Sequential patterns of colonization of coarse woody debris by Ips pini (Say) (Coleoptera: Scolytidae) following a major ice storm in Ontario

Abstract

• 1 It is widely known that many bark and wood‐boring beetle species use nonresistant coarse woody debris (CWD) created by disturbances; however, the ability of these secondary species to cause mortality in healthy trees following a build‐up of their populations remains unclear. We characterized the pattern of colonization by Ips pini (Say) following a major ice storm that created large amounts of CWD varying in resistance to colonization (i.e. ranging from snapped tops with no resistance to heavily damaged trees with intact root systems). A major question was how the beetles responded to the different types of storm‐damaged material and whether healthy undamaged trees were colonized and killed following increases in beetle populations.
• 2 Six red pine, Pinus resinosa Ait., plantations in eastern Ontario were monitored from 1998 to 2001 inclusive: three with high storm damage (approximately 120 m³/ha CWD) and three with minimal damage (approximately 20 m³/ha CWD). Transects (200 × 2 m) were sampled yearly in each plantation to assess the type and amount of damaged pine brood material colonized by the pine engraver beetle, I. pini.
• 3 Beetles preferentially infested the most nonresistant material available each year (i.e. all snapped tops in year 1, all standing snags, up‐rooted trees and many bent trees by year 2, but still less than 50% of trees blown over but with intact root systems by year 3). By years 3 and 4, the majority (approximately 75%) of severely damaged trees (with > 50% crown loss) died prior to beetle colonization.
• 4 The size of the beetle population tracked the abundance of available woody material from year‐to‐year within a plantation; populations were very large in the first 2 years, and declined significantly in the last 2 years.
• 5 Healthy standing red pines were apparently resistant to colonization by the beetles, despite the significant build‐up in their populations. Hence, the results of the present study suggest that native bark beetle populations will not cause further tree mortality following such a disturbance in this region.

     

Tree species and position matter: the role of pests for survival of other insects

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Histochemical and biochemical approaches to the study of phenolic compounds and peroxidases in needles of Norway spruce (Picea abies)

The three youngest age-classes of needles of Norway spruce ( Picea abies ) were collected from four sites in the Krusne Hory Mountains (Czech Republic) characterized by different levels of damage caused by environmental pollution. Histochemical methods did not reveal any differences in localization of phenolics among the needles. Mesophyll cells close to the epidermis of needles and cells around resin ducts and substomatal cavities often accumulated higher amounts of phenolics than the rest of the mesophyll cells, but this was independent of age and damage. Needles of different age- and damage-class did not show any marked changes in general lignification pattern. However, a lower intensity of histochemical detection of lignin was observed in needles from the most damaged site. This finding was confirmed by chemical analysis using thioglycolic acid. Generally, the amount of lignin in mesophyll cells was lower in damaged trees than in healthy ones. Using the Folin–Ciocalteau method, no significant differences in the total content of phenolics were observed in the needles, although HPLC revealed marked alterations in the forms of seven phenolic acids. Concentrations of conjugated forms of phenolic acids (esters and glycosides) were higher in damaged needles (255.9 μg g⁻¹ f. wt) than in healthy needles (189.8 μg g⁻¹ f. wt). By contrast, content of esterified phenolic acids incorporated into cell walls was higher in needles from healthy trees (101.1 μg g⁻¹ f. wt) than in damaged needles (78.3 μg g⁻¹ f. wt). Marked differences were also observed in the activity of soluble peroxidases, although the activity of ionically bound forms was approximately the same in healthy and damaged needles. The total amounts of chlorophylls and carotenoids decreased as environmental damage increased.     

Diagnostic parameters for selecting against novel spruce (Picea abies) decline: I. Tree morphology and photosynthesis response to acute SO2 exposures

The dose- and time-response effects of single 4-h day-exposures to 0.50, 0.79, 1.28, 1.58, 2.38 or 3.35 μl l^?1 (ppm) SO₂ followed by single 3-h night-exposures of 0.60, 0.87, 1.54, 1.91, 2.91 or 3.98 μl l^?1 SO₂ on photosynthesis, transpiration and dark respiration were examined for nine East European (Carpatho-Ukrainian, ‘Rachovo’) half-sib families and for two populations, one from the FRG (‘Westerhof’) and one from the GDR (‘Schmiedefeld’) of Norway spruce [Picea abies (L.) Karst.], all in their 4th growing season. Even the lowest SO₂ concentration reduced photosynthesis and transpiration within 1 h. Photosynthesis of the different spruce types was affected significantly differently, the most sensitive spruce being suppressed 2.5 times more than the most tolerant spruce. ‘Westerhof’ was more resistant to SO₂ than the average ‘Rachovo’ half-sibs. Neither transpiration (stomatal reaction), which was affected alike by all SO₂ concentrations, nor SO₂ uptake, explained adequately the effects on photosynthesis. Night transpiration, but not dark respiratin, was stimulated by night SO₂ preceded by day SO₂ exposure. The gradient of different SO₂ sensitivities among young trees from the half-sib families demonstrated a significant negative correlation with the gradient of different sensitivities to novel decline symptoms of their parents growing in a rural seed orchard in Denmark, and with the gradients of four morphology parameters, (height, branching, branch density and the number of Lammas shoots) of the young trees, which in turn demonstrated a positive correlation with decline sensitivity in the seed orchard. The relative photosynthesis sensitivity and the morphology of half-sibs may serve as diagnostic parameters for laboratory selection of the most resistent trees to novel spruce decline in the field. There was a positive correlation between SO₂ induced scorching of Lammas shoots and the inhibition of photosynthesis, but not between the severity of SO₂ scorching and symptoms of novel spruce decline. The two visible types of symptoms looked very different.