Fungi isolated from Picea abies infested by the bark beetle Ips typographus in the Białowieża forest in north‐eastern Poland

The assemblage of fungi occurring in the sapwood of Norway spruce (Picea abies) and in bark beetle galleries following attack by the Eurasian spruce bark beetle Ips typographus was investigated in the Bia?owie?a forest in north‐eastern Poland. Fungi were isolated from blue‐stained sapwood of beetle‐infested spruce trees in June 2002, and a few isolates were also obtained from ascospores and conidia taken from perithecia and asexual structures occurring in the gallery systems of the insects. The mycobiota of I. typographus in the Bia?owie?a forest was dominated by ophiostomatoid fungi, which were represented by seven species. Four species, including Ceratocystis polonica, Grosmannia penicillata, Ophiostoma ainoae and Ophiostoma bicolor were isolated at high frequencies, whereas three other taxa, Ceratocystiopsis minuta, Ceratocystiopsis alba and a Pesotum sp. were rare. The anamorphic fungus Graphium fimbriisporum and yeasts also occurred occasionally. In addition, the basidiomycete Gloeocystidium ipidophilum was relatively common. The pathogenic blue‐stain fungus C. polonica was the dominant fungal associate of I. typographus in the Bia?owie?a forest, which is consistent with a previous study at this area in the 1930s. Ceratocystis polonica was the most frequently isolated species at the leading edge of fungal colonization in the sapwood and had on an average penetrated deeper into the wood than other fungal associates. This suggests that it acts as a primary invader into the sapwood after attack by I. typographus in the Bia?owie?a forest, followed by O. bicolor, O. ainoae, G. ipidophilum and G. penicillata. Thus far, the Bia?owie?a forest is one of the few areas in Europe, where C. polonica has been reported as a dominate fungal associate of I. typographus.     

Ophiostomatoid fungi associated with bark beetles onAbies veitchii in wave-regenerated forests

Relationships between tree mortality and bark beetle infestation onAbies veitchii at the wave-regenerated forest in Mt. Asahi, Okuchichibu area were investigated. Most of the firs with green needles and newly developed current year’s shoots in the dieback zone were heavily infested by bark beetles before the death of the trees. After heavy infestation of beetles, about half of the infested firs died within the year, and the other half died in the next year. When the species composition of bark beetles and associated ophiostomatoid fungi were investigated in Mt. Asahi and also at a typical wave-regenerated forest in Mt. Shimagare, Yatsugatake area,Cryphalus montanus andC. piceae were dominant beetle species for Mt. Asahi and Mt. Shimagare, respectively.Ophiostoma subalpinum andO. europhioides were dominant fungal species at both wave-generated forests. BecauseO. subalpinum was detected more frequently from deeper areas of sapwood thanO. europhioides, it was suggested that the fungal species may accelerate the death of stressed firs in wave-regenerated forests. Contribution No. 169, Laboratory of Plant Parasitic Mycology, Institute of Agriculture and Forestry, University of Tsukuba.     

Occurrence of pathogens in associated living bark beetles (Col., Scolytidae) from different spruce stands in Austria

Pathogen occurrence was studied in 16?099 adult specimens of 10 different bark beetle species, which live associated on Norway spruce (Picea abies (L.) Karst.). Beetles (mainly Ips typographus L. and Pityogenes chalcographus L.) were collected from 6 different localities in Austria (4 secondary spruce stands and 2 natural forest type stands, 9 sampling plots in total) in elevations between 400?m and 1600?m. Various viral, protozoan, and fungal pathogens could be diagnosed with a light microscope in the examined beetles. Numerous pathogen species were known from former studies, some pathogens were totally new or could be found in a new host species beside their type host. The most dominant pathogen species were Protozoa, Gregarina cf. typographi, Malamoeba cf. scolyti, and Chytridiopsis cf. typographi. Over the whole investigation period, the highest pathogen diversity with eight pathogen species was found in I. typographus. Differences were observed in the pathogen complex of each beetle species from the different collection sites and in different years of investigation. Several species showed an overlapping in their host range and infected various bark beetle species. Furthermore, pathogen occurrence and prevalence differed in bark beetles from 4 different sampling plots in an area (one locality) within a distance of a few kilometres.     

Ophiostomatoid fungi associated with root‐feeding bark beetles on Scots pine in Poland

Ophiostomatoid fungi are known to be associated with various species of bark beetles. However, information about fungal associates of root‐feeding bark beetles in Europe is still fragmentary. For this reason, the fungal associates of Hylastes ater, H. opacus and Hylurgus ligniperda on Pinus sylvestris were isolated and identified. A total of 743 fungal isolates were collected and separated into 10 morphological groups. Analyses of ITS rDNA and partial β‐tubulin gene sequences confirmed that these groups represented distinct species. The 10 species included a total of 13 associations between fungi and bark beetles that had not been recorded previously. All of the bark beetles examined were frequently associated with ophiostomatoid fungi. The fungal diversity and relative abundance of species were very similar in the three species of root‐feeding bark beetles. The most commonly encountered associates of these beetles were Grosmannia radiaticola, Leptographium lundbergii, L. procerum and L. truncatum. Insect infestation data furthermore suggest that Hylastes spp. and Hg. ligniperda are also important vectors of the fungal pathogen Sphaeropsis sapinea.     

Diversity of Ophiostomatales species associated with conifer-infesting beetles in the Western Carpathians

Bark beetles (Coleoptera: Curculionidae, Scolytinae) are commonly recognised as important agents of tree mortality in coniferous forests of the Western Carpathians. They, together with weevils, are consistently associated with ophiostomatoid fungi. Information regarding conifer beetle-associated fungi in the Western Carpathians remains incomplete and unreliable, particularly with respect to fir-infesting bark beetles. This study aims to clarify associations between fungi in the genera Graphilbum, Leptographium, Ophiostoma and Sporothrix (Ophiostomatales) and their beetle vectors in Norway spruce (Picea abies), European larch (Larix decidua) and silver fir (Abies alba). Samples associated with 20 bark beetle species and weevils were collected from nine stands in Poland and the Czech Republic. Fungi were isolated from adult beetles and galleries. Isolates were identified based on morphology, DNA sequence comparisons for four gene regions (ITS, LSU, ß-tubulin, TEF 1-α) and phylogenetic analyses. In total, 46 distinct taxa were identified, including 25 known and 21 currently unknown species. Several associations between fungi and subcortical insects were recorded for the first time. In addition, O. borealis and O. quercus were detected from A. alba for the first time. The composition of the fungal communities varied among the studied tree species and to a lesser degree among the beetle species. The spruce-infesting bark beetles were commonly associated with species of Leptographium s. l. and Ophiostoma s. str.; the larch-infesting bark beetles were often associated with Ophiostoma s. str. and Sporothrix, while the fir-infesting bark beetles were commonly associated with Ophiostoma s. str. and Graphilbum. The most commonly encountered fungal associates of the examined insects were (a) Grosmannia cucullata, G. piceiperda, Grosmannia sp. 1, Ophiostoma macroclavatum and O. piceae with the spruce-infesting bark beetles; (b) O. pseudocatenulatum and Sporothrix sp. 1 with the larch-infesting bark beetles; and (c) O. piceae, Ophiostoma sp. 2 and Graphilbum sp. 2 with the fir-infesting bark beetles. The differences in fungal associates among the bark beetle species occurring on P. abies, L. decidua and A. alba could be linked to the different habitats that these beetles occupy.     

Ophiostomatoid fungi associated with three spruce-infesting bark beetles in Slovenia

Context

Ophiostomatoid fungi can severely affect the health and economic value of Norway spruce trees (Picea abies). Although the diversity of ophiostomatoid species and their associations with insects have been well-investigated in central and northern Europe, little is known about the conditions in south-eastern Europe.

Aim

This study aims to study the assemblages of ophiostomatoid fungi associated with three bark beetle species (Ips typographus, Ips amitinus, and Pityogenes chalcographus) that infect Norway spruce in Slovenia.

Methods

Bark beetles were sampled in four phytogeographic regions in Slovenia. The fungi found on the bark beetles were identified based on morphology, DNA sequence comparisons of ITS regions and phylogenetic analysis. The species compositions of the fungal associates of the three insect species were compared and the pairwise associations of the occurrence of the fungal species were analysed.

Results

Thirteen different species were found. The most commonly encountered fungal associates of the beetles were Ophiostoma bicolor, Ophiostoma brunneo-ciliatum, Grosmannia piceiperda, Ophiostoma ainoae, Ceratocystiopsis minuta, and Grosmannia penicillata. The composition of the fungal associates differed among the bark beetle species, but not among the phytogeographic regions.

Conclusions

This study confirms that ophiostomatoid species are common associates of the investigated bark beetle species. Many ophiostomatoid species have strong host associations. I. typographus and P. chalcographus can act as effective vectors for O. bicolor, O. ainoae, G. piceiperda and O. brunneo-ciliatum, whereas I. amitinus often carries G. piceiperda and C. minuta in Slovenian forests.     

Intraspecific variation in Heterobasidion annosum for growth in sapwood of Picea abies and Pinus sylvestris

Two greenhouse experiments were conducted to study intraspecific variation in growth of the root rot fungus Heterobasidion annosum in living host sapwood. In experiment 1, Scots pine (Pinus sylvestris) and Norway spruce (Picea abies) seedlings were inoculated with H. annosum isolates, 14 each of the S-and P-intersterility groups, collected from various parts of Sweden. In pine, the P-group isolates were more virulent than the S-group isolates both in terms of infection frequency, induced mortality rate (p < 0.05), and fungal growth in sapwood (p < 0.05). In spruce, the P-group isolates were also more virulent on average, but the difference was not statistically significant. Both S and P isolates had a higher infection frequency and a significantly longer sapwood growth on spruce than on pine. The P-group caused higher mortality on pine than on spruce. The length of the lesion in the inner bark was strongly correlated with fungal growth in spruce, but not in pine where the lesions were short or absent. In experiment 2, ten Norway spruce clones were inoculated with 18 S-isolates, originating from nine live-decayed trees and from nine spore-infected stumps in a single Norway spruce stand. The objective was to test whether any selection for growth rate in sapwood was detectable among individuals of H. annosum originating either from stumps or trees. The results gave no support for such selection since no difference in sapwood growth between the two groups of isolates was found.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Fungi associated with Ips typographus on Picea abies in southern Poland and their succession into the phloem and sapwood of beetle‐infested trees and logs

This study dealt with the species distribution and frequency of fungi associated with the bark beetle Ips typographus (Scolytidae) on spruce trees of various states of health in southern Poland. The spruce trees were assessed by their degree of defoliation and damage of their crowns and trunks after attack by I. typographus. The state of health of trees from which samples were obtained was related to varying stages of brood development of I. typographus. Fungi were isolated from phloem taken from and around insect galleries and the sapwood underneath brood systems. Samples were taken from ‘healthy‐looking’, weakened, wind‐fallen and wind‐broken trees as well as from trap trees. The mycobiota associated with I. typographus was quite diverse in respect of the number of detected species, 65 fungal taxa were obtained from the phloem of trees infested by I. typographus, and 36 taxa occurred in the sapwood underneath insect galleries. The spectrum of fungi mainly consisted of ascomycetes and anamorphic fungi. The ophiostomatoid fungi were represented by 14 species and were the most numerously represented group in all niches examined. The most frequent ophiostomatoid species were Ceratocystis polonica, Ophiostoma ainoae, O. bicolor, O. penicillatum, O. piceae and O. piceaperdum. The frequency of occurrence of ophiostomatoid fungi differed significantly between the phloem and sapwood as well as in relation to the varying states of health of the spruce trees. These quantitative differences in the mycobiota of I. typographus between spruce trees belonging to different health categories can be explained by successional patterns of fungal colonization of host tissues following attack by I. typographus. The pathogenic species C. polonica was the primary invader, occurring most frequently in the sapwood of ‘healthy‐looking’ trees. Ophiostoma bicolor, O. penicillatum and O. piceaperdum also occurred during the early stages of brood development of I. typographus on ‘healthy‐looking’ trees, but they mainly colonized the phloem. In contrast, O. ainoae, O. minuta and O. piceae likely follow the aforementioned species as secondary and tertiary invaders into the phloem and the sapwood of spruce trees.     

Fungal succession in sapwood of Norway spruce infested by the bark beetle Ips typographus

Fungal invasion of Norway spruce sapwood was investigated during a period of 130 weeks fol-lowing attack by the spruce bark beetle Ips typographus. Heavy beetle attack tended to overwhelm the trees quickly and no secondary resinosis was produced. Nonetheless, fungal penetration in the sapwood was initiated only slowly, although it accelerated during the fourth week after attack. Typically the leading edge of fungal penetration was a few millimeters in advance of the visible blue-stain during the sapwood colonization. Fungi invaded the sapwood in a successional pattern with the most pathogenic species Ophiostoma polonicum first, followed by other beetle-transmitted Ophiostoma and Graphium species. The later succession was probably influenced by sapwood moisture. Strong decaying white-rotters invaded near the base of the trees, where sapwood moisture remained favourable, while cf. Trichoderma viride dominated in the drier parts or the trees.     

Effects of forest restoration treatments on the abundance of bark beetles in Norway spruce forests of southern Finland

Restoration of protected areas in boreal forests frequently includes creating substantial volumes of dead wood. While this benefits a wide range of dead wood dependent invertebrate species, some of these are regarded as forest pests. Therefore, the risk of elevated levels of tree mortality in surrounding commercial forests must be considered. In a large-scale field experiment in southern Finland, we studied the effects of restoration treatments on the abundance of bark beetles within and in the vicinity of restored areas, in particular focusing on Ips typographus and Pityogenes chalcographus. The treatments applied to managed Norway spruce forests were controlled burning and partial harvesting combined with retaining 5, 30 or 60 m³/ha of cut down wood. We found that the abundance of bark beetles increased by both burning and harvesting with down wood retention, being highest where burning and harvesting had been combined. The actual volume of down wood retention had no significant effect. The effect of burning on the number of bark beetles along host tree boles was negative which suggests that burnt spruces provided a less suitable resource for bark beetles than unburnt dead spruces. The abundance of bark beetles along host trees also decreased with increasing volume of down wood retention. The abundance of P. chalographus was slightly elevated up to 50 m outside restored areas but the abundance was very low compared to that within the areas. The abundance of I. typographus was extremely low outside restored areas. We conclude that restoration treatments increase the abundance of bark beetles via increased availability of resources, but that the effect of burning is likely to be counteracted by decreased resource quality. Thus, burning might be the “safest” way to produce large quantities of dead wood. Furthermore, the fact that only few beetles were collected in adjacent areas suggests that restored areas pose little threat of serving as refugia in which bark beetle populations increase in sufficient numbers to attack live trees in adjacent forests. However, restoration actions repeated at consecutive years within a small area might enable the populations to grow to outbreak levels.     

Variation in growth of Heterobasidion parviporum in a full-sib family of Picea abies

Abstract

Heterobasidion parviporum (Fr.) Niemelä & Korhonen and Heterobasidion annosum (Fr.) Bref. sensu lato are some of the major forest pathogens in the northern hemisphere causing root and butt rot to conifers. The relative susceptibility to H. parviporum was investigated in a full-sib family of Norway spruce [Picea abies (L.) Karst.] by inoculating a set of 252 cloned progenies from a controlled cross. Four ramets of each progeny were used and the 2-year-old rooted cuttings were incubated for 6 weeks under greenhouse conditions. The condition of the cuttings was assessed visually and all the plants were in excellent vigour with no mortality recorded during the experiment. To score the relative susceptibility, lesion length in the inner bark and fungal growth in the sapwood were measured. Among the progenies, significant differences were found for fungal growth in the sapwood (p<0.0005). There was no significant difference for lesion length; however, there was a significant positive correlation between fungal growth and lesion length. The broad-sense heritability was 0.11 for fungal growth. This shows that the genetic component for susceptibility to H. parviporum can be detected even within a full-sib family of Norway spruce and that there is a potential for mapping quantitative trait loci for this trait in Norway spruce.     

Induced resistance of Norway spruce,variation of phenolic compounds and their effects on fungal pathogens

Three clones of Norway spruce (Picea abies) were studied for their response to mass‐inoculation with the blue‐stain fungus Ceratocystis polonica. The effect of different pretreatments (fungal inoculation and wounding) before mass‐inoculation was investigated for their possible role in an acquired resistance reaction. Pretreated trees showed enhanced resistance to the subsequent mass‐inoculation relative to control trees that received no pretreatment. Furthermore, the fungal colonization of inoculated trees was less than that of wounded trees. The phenolic content of the bark, analysed by RP‐HPLC, was compared in trees receiving different treatments. Trees inoculated with C. polonica had higher average concentration of (+)‐catechin, taxifolin and trans‐resveratrol than wounded trees. Both inoculated and wounded trees had higher average concentrations of these compounds than control trees. The effect of the phenolic extract of Norway spruce bark on the growth of the root rot fungus Heterobasidion annosum and the blue‐stain fungi C. polonica and Ophiostoma penicillatum were investigated in vitro. Heterobasidion annosum was not negatively affected, and the extracts had fungistatic effects on the blue‐stain fungi. The growth of O. penicillatum was more inhibited than the growth of the more aggressive C. polonica.     

Association of the pine-infesting Pissodes species with ophiostomatoid fungi in Poland

Bark weevils are consistently associated with various fungi. They act as effective vectors for root-rot fungi, the rust pathogen and the ophiostomatoid fungi. In comparison with bark beetles, the interaction between ophiostomatoid fungi and bark weevils has been poorly studied in Europe. This study aims to clarify the ties among ophiostomatoid fungi and their weevil vectors in Pinus sylvestris. Samples associated with three bark weevils, including Pissodes castaneus, P. piniphilus and P. pini were collected from seven pine stands in Poland. Fungi were isolated from laboratory-reared and field-collected beetles, larvae and galleries of weevils. Isolates were identified based on morphology, DNA sequence comparison for two gene regions (ITS, ß-tubulin) and phylogenetic analyses. Fourteen morphological and phylogenetic ophiostomatoid species were identified among the 1,219 isolates. These 14 species included 11 novel associations between fungi and bark weevils. The most commonly encountered fungal associates of bark Pissodes species were Leptographium procerum, Ophiostoma quercus and O. minus. The spectrum of fungal associates was similar in three bark weevil species, despite some differences between species. Except for P. castaneus, all species of bark weevil were frequently associated with ophiostomatoid fungi. Pissodes castaneus does appear to be a consistent vector of L. procerum and Sporothrix inflata while O. minus, O. quercus and O. cf. rectangulosporium were often found in association with P. piniphilus and P. pini.     

Analysis of Norway spruce dieback phenomenon in Latvia – a belowground perspective

In 2010, dieback of Norway spruce (Picea abies) was observed in Latvia. As mostly stands on seasonally or permanently water-saturated soils were damaged, we hypothesized that the recorded foliar damage might be associated with belowground factors. We established 48 sample plots in stands with damage symptoms for soil parameter evaluation and fine root sampling to determine the number of fine roots in different growth stages and to describe the associated fungal community. Among sample plots on organic soils, there were fewer viable older fine roots and greater number of dead fine roots in more damaged sites than in less damaged. These root parameters were significantly correlated with higher groundwater level and soil pH. The fungal community was dominated by ectomycorrhizal (ECM) species (the most common was Tylospora asterophora), the endophyte Oidiodendron maius and saprotrophic species Cryptococcus magnus. Saprotrophic species had higher abundance in more damaged sites, suggesting a shift in fungal communities from ECM fungi. In conclusion, the results of this work suggest Norway spruce root system response to short-term climatic stress, which should be considered when planning spruce forest management, especially on organic soils with a high groundwater level.     

Stimulation of tree defenses by Ophiostomatoid fungi can explain attack success of bark beetles on conifers

? Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.

? After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.

? We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.

1. Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.
2. We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.

? We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).

? Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:

1. any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;
2. the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;
3. Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.

? We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.

? We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.

     

Colonisation of native and exotic conifers by indigenous bark beetles (Coleoptera: Scolytinae) in France

Planting exotic conifers offers indigenous forest insects an opportunity to extend their host range and eventually to become significant pests. Knowing the ecological and evolutionary modalities driving the colonisation of exotic tree species by indigenous insects is thus of primary importance. We compared the bark beetle communities (Coleoptera: Curculionidae, Scolytinae) associated with both native and introduced conifers in France. The aim of our study was to estimate the influence of both host- and insect-related factors on the beetles’ likelihood to shift onto new hosts. We considered the influence of host origin (i.e. native vs. exotic), host tree species identity, tree bark thickness and tree taxonomic proximity, as well as insects’ host specificity. A field inventory using trap trees was carried out in two regions in France (Limousin and Jura) during two consecutive years (2006 and 2007) on three European native conifer species [Norway spruce (Picea abies); Scots pine (Pinus sylvestris) and European Silver-fir (Abies alba)] and five North American [Sitka spruce (Picea sitchensis); Eastern white pine (Pinus strobus); Grand fir (Abies grandis); Douglas fir (Pseudotsuga menziesii) and Western red cedar (Thuja plicata)]. A total of 18 indigenous and 2 exotic bark beetle species were collected. All exotic conifer species were colonised by indigenous bark beetle species and no significant difference was observed of the cumulated species richness of the latter between native and exotic tree species (13 vs. 14, P < 0.05). The ability of indigenous bark beetles to shift onto exotic conifers appeared to strongly depend on host species (significantly structuring bark beetle assemblages), the presence of phylogenetically related native conifer species and that of similar resources, in combination with insect host specificity. Host tree species status (native or exotic) also seemed to be involved, but its effect did not seem as essential as that of the previous factors. These findings are discussed in terms of adaptation, plasticity and practical aspects of forest management.     

Comparative susceptibility of pine,spruce and larch to sapstain

Summary This study compared the susceptibility of five UK‐grown conifer species to colonization by sapstain fungi in two trials carried out in consecutive years. The conifers consisted of Sitka spruce (Picea sitchensis), Japanese larch (Larix kaempferi), Norway spruce (Picea abies), Scots pine (Pinus sylvestris) and lodgepole pine (Pinus contorta). Freshly cut 1‐m logs were exposed to the available inoculum of sapstain fungi from April to August in a woodland environment in the south east of England. Logs of each species were removed after 1‐, 2‐ and 4‐month exposure and sampled destructively to assess the amount of sapstain. In the second trial, per cent moisture content and concentrations of nitrogen, carbohydrate and phenolic compounds in the sapwood were also measured at the start and end of the trial. After 2 months, only the sapwood of both pine species had significant levels of sapstain; mean values of 37% and 19% for lodgepole pine (year 1 and year 2 respectively) and 12% and 1% for Scots pine. After 4 months, the levels of sapstain in both pine species exceeded 60% in both years. By contrast, very little sapstain developed in the other conifer species with maximum mean values of 10% for Norway spruce, 3.5% for larch and less than 1% for Sitka spruce. Overall, the moisture content of the logs decreased progressively in all species over the length of the trial. However, pine logs tended to retain higher levels of moisture throughout the trial compared with spruce or larch. The relatively higher moisture content of pine sapwood may be closer to the optimal moisture content that sapstain fungi require for infection and colonization, thereby contributing to the increased susceptibility of pine compared with the other conifer species. The pine logs also suffered from some colonization by bark beetles (Ips sexdentatus), which increased the inoculum potential and the opportunity for colonization by sapstain fungi. In addition, particular phenolic compounds in conifer sapwood may play a role in determining the resistance of some species to sapstain. Notably the most resistant species, Sitka spruce, was the only softwood that still retained detectable levels of phenolics in the sapwood to the end of the trial.     

The virulence of phytopathogenic fungi associated with the bark beetles Tomicus piniperda and Orthotomicus erosus in Tunisia

Phytopathogenic fungi associated with the bark beetles Tomicus piniperda and Orthotomicus erosus were isolated in various pine forests of Tunisia. Tomicus piniperda and its galleries yielded Leptographium wingfieldii, Ophiostoma minus, and Ophiostoma ips. Ophiostoma minus was the most frequent species associated with T. piniperda, in both the attacking and the emerging beetles. It was collected from most investigated forests, whereas O. ips and L. wingfieldii were obtained only from forests located in Central and Northern Tunisia. Frequencies of association with T. piniperda were always low, reaching 11.1% only once, for O. ips. Ophiostoma ips was the only blue stain fungus associated with O. erosus and its galleries. It was found in all the localities, but at a low and variable frequency, exceeding 15% very rarely. The virulence of 16 fungal isolates was tested by single inoculations into Pinus halepensis (Aleppo pine) at two localities. Differences were detected among species; L. wingfieldii was the most virulent and O. minus the least virulent species in terms of phloem reaction zone formation and fungal growth in the phloem. In a separate experiment, mass inoculations (400 and 800 inoculations per m² of bole on 1‐m high belts) were performed with two isolates of L. wingfieldii on Aleppo pine and on Pinus brutia (Brutia pine). Three months later, measurements of sapwood status (% of conductive transversal section) and of its specific hydraulic conductivity, as well as of the phloem reaction zone lengths, did not show any isolate or density effect. At these experimental sites, Brutia pines appeared significantly more susceptible than Aleppo pines.     

Associations between Pityogenes bidentatus and fungi in young managed Scots pine stands in Poland

The association between Pityogenes bidentatus and fungi was studied in young, managed Pinus sylvestris stands in Poland. Fungi were isolated from emerged adults and their galleries collected from four populations. In total, 2089 fungal isolates including 42 species, were obtained. Penicillium sp. 1 and Geosmithia sp. 1 were the most commonly isolated fungi from beetles (49% and 41% of beetles respectively). Geosmithia sp. 1 species was the dominant species in P. bidentatus galleries with a frequency of occurrence of 57.9%. Hormonema dematioides was the second most abundant fungus in gallery systems (17.1% of wood samples). Two of the isolated Geosmithia species were previously undescribed. Pityogenes bidentatus also vectored three ophiostomatoid species: Ophiostoma minus, O. piceae and Graphium sp. ‘W’. These species were occasionally isolated from beetles and their galleries, suggesting a non-specific relationship.