Occurrence,genetic diversity,and potential pathways of entry of Halyomorpha halys in newly invaded areas of Canada and Switzerland

Halyomorpha halys has recently established in countries outside of its Asian area of origin. In the North Eastern USA, this pest causes severe economic loss in agricultural crops. Breeding populations have been confirmed in Switzerland and Canada; however, their pathways of entry and genetic diversity remain unknown. To determine invasion pathways and source population(s), the diversity of specimens from Asia, North America, and Europe was investigated. Genetic diversity in Asia was higher than that in the Canada and Switzerland. Although three haplotypes were detected among Canadian H. halys, one haplotype dominated. In contrast, two out of three haplotypes were found frequently among Swiss samples. Based on molecular data and interception records it appears likely that H. halys in Canada is derived from the movement of established US populations. Further, North American populations likely originated from the Hebei/Beijing regions of China. The area of origin of the Swiss samples remains unclear. Although the dominant haplotype in Switzerland was consistent with Asian samples collected in the Hebei and Beijing provinces, it was not the dominant haplotype in these regions and further sampling is necessary to its confirm the distribution and abundance in the area of origin. The remaining two haplotypes were unique to Switzerland and no matches were found among our Asian samples. Interestingly, no haplotypes were shared between the North American and Swiss samples, indicating that the two invasions were separate and distinct events and did not result from the movement of goods and materials between USA and Europe.     

The North American P group of Heterobasidion annosum s.l. is widely distributed in Pinus pinea forests of the western coast of central Italy

The distribution of the North American P group of Heterobasidion annosum s.l., recently reported from a Pinus pinea forest in the surroundings of Rome, was studied using mating tests and DNA fingerprinting (mitochondrial DNA, random amplified microsatellite technique and two group‐specific markers). This fungus is present in several forests and small plantations along the Tyrrhenian coast of the Italian peninsula, within an area approximately 100 km long, extending from Fregene in the north to the National Park of Circeo in the south, and 27 km wide including the city of Rome. In pine forests of Castelporziano, Castel Fusano and Anzio, where US troops resided during the Second World War, the North American P group is more frequent than the European P group. The low number of mating alleles in the Italian population of the North American P group supports the hypothesis of its origin from a small number of introductions. The near 100% sexual compatibility between the North American and European P groups, together with inconsistencies in results obtained with different identification methods of these groups, suggests that hybridization between the North American and European P populations occurs occasionally.     

Heterobasidion annosum group S identified in north-eastern China

Samples of Heterobasidion annosum were collected in two stands of mixed forest in Changbaishan Nature Reserve in north-eastern China. Pure cultures were isolated from 15 specimens growing on Abies, Picea, Pinus and Populus. Mating tests showed that all these collections belong to the S inter-sterility group. The compatibility of Chinese homokaryons with S homokaryons from Europe was 100%. Clamp connections appeared almost equally in both strains of each pairing, and no distinct demarcation line formed in the pairings. The mating frequency of Chinese isolates with the European F group was also high, approximately 94%. These matings, however, were usually unilateral and a more or less distinct demarcation line was present in most of the pairings. The Chinese isolates were completely intersterile with the European P group. The S group of H. annosum in north-eastern China appears to have similar morphology and ecology as in Europe. It is common on Abies and Picea, but is not a serious pathogen in the pristine forests of north-eastern China.     

Larval performances and life cycle completion of the Siberian moth, <Emphasis Type="Italic">Dendrolimus sibiricus</Emphasis> (Lepidoptera: Lasiocampidae), on potential host plants in Europe: a laboratory study on potted trees

The Siberian moth, Dendrolimus sibiricus, Tschtv. is the most harmful defoliator of coniferous forests in North Asia. The pest has already spread over the Urals and continues moving westwards. Recently, it has been recommended for quarantine in member countries by European and Mediterranean Plant Protection Organization (EPPO). The performances of the pest on coniferous species planted in Europe were assessed on a range of potted trees corresponding to the spectrum of economically important conifers in the EU: European larch Larix decidua, Norway spruce Picea abies, Scots pine Pinus sylvestris, European black pine Pinus nigra, and the North American species: Douglas fir Pseudotsuga menziesii and grand fir Abies grandis. Larvae showed a potential to survive and complete the development on all these host tree species. Favorable hosts were grand fir, European larch, and Douglas fir that allowed higher survival, better larval development, and as a result, yielded heavier pupae and adult moths with higher longevity. Black pine was a poor host but, however, could still support larval and pupal development. Norway spruce and Scots pine had an intermediate behavior. If accidentally introduced to Europe, the Siberian moth may become especially damaging in forest stands predominated by European larch and by the North American firs. Norway spruce and especially the two-needle pines will be less prone to intensive defoliation by this species. The fact that the pest may damage the range of economically important coniferous species should be taken into account in the pest risk assessment for Europe and also for North America where the Siberian moth occurrence is considered likely.     

Biological control of Heterobasidion annosum on Pinus pinea by Phlebiopsis gigantea

The efficacy of a native isolate of Phlebiopsis gigantea in protecting the stumps of Pinus pinea against spore infection by Heterobasidion annosum s.str. was investigated. In preliminary tests carried out in stem pieces of P. pinea in confined environment, the efficacy of the isolate was compared with Rotstop^®, the commercial formulation prepared from a north European P. gigantea strain. Both showed a fully protective effect against artificial inoculation of H. annosum. Moreover, in stump treatment experiments carried out in a coastal P. pinea forest close to Rome, the native P. gigantea was effective against heavy airborne inoculum of H. annosum. Some natural infection by P. gigantea occurred in the stumps, but it was unable to control the pathogen. Random amplified microsatellite analysis allowed to recognize the presence of different P. gigantea strains in the forest.     

Investigations on Heterobasidion annosum s.lat. in central and eastern Asia with the aid of mating tests and DNA fingerprinting

To investigate the taxonomy of Heterobasidion in Eurasia, 49 specimens belonging to H. annosum sensu lato from Asia were identified with the aid of mating tests. Most of the specimens originated from north‐eastern and south‐western China and from the Altai region in southern Siberia, but a few isolates from Kirghizia, Japan and India were also tested. In addition to mating tests, the material from China was investigated with DNA fingerprinting. Heterobasidion annosum sensu stricto was identified only from the Altai region. Homokaryotic isolates from other specimens, except the Indian ones, were sexually compatible with H. parviporum, but they also showed a high degree of compatibility with H. abietinum and with the North American S group. The isolates from SW China (eastern Himalayas) mated with about equal frequency with the European strains of H. parviporum and H. abietinum. However, the DNA fingerprinting showed that these isolates were more closely related to H. parviporum, and hence they were tentatively included in this species. The North American S group was more distant from these Eurasian taxa. Four old isolates from India mated only weakly with the members of the H. annosum s.lat. According to the species concept presented, the distribution of H. parviporum extends from western Europe through southern Siberia to northern China, Japan and the eastern Himalayas. H. annosum s.str. is so far identified only from the Altai region outside Europe, and H. abietinum only from Europe.     

Fungi dangerous at Pinus contorta with special reference to pathogens from North Europe

Examples are given of North American fungi potentially dangerous to Pinus contorta plantations in northern Europe. The pathogenicity of North European fungi is discussed. P. contorta was found to be immune or nearly immune to all European rust fungi, more resistant than P. sylvestris to Phacidium infestans and Lophodermium pinastri, but less resistant to Crumenulopsis sororia and Discella strobilina. For other fungi no such clearcut conclusions could be drawn.     

Unbemerkte Einbürgerung und Ausbreitung des nordamerikanischen Grubenhalsigen Splintholzkäfers,Lyctus cavicollis LeConte,in Mitteleuropa,nebst Anmerkungen zur möglichen Einschleppung zweier anderer nearktischer Lyctiden (Coleoptera,Lyctidae)

Undiscovered introduction and distribution of the North American Western Powder-post BeetleLyctus cavicollis LeConte in Central Europe, with remarks to possible importations of two other nearctic lyctids (Coleoptera; Lyctidae)The article analyses 30 reports on the occurrence ofLyctus cavicollis LeC. introduced from North America into southwest-central Europe since 1974 and considers the circumstances of the finds. 18 synanthropic occurrences are known, along with ten finds in the wild, at least three of which were in nearly natural established populations. BecauseL. cavicollis is already widely distributed and established in parts of Central Europe, it is impossible today to reconstruct even approximately the chronological history of its introduction.On the other hand, only two cases are known from the last forty years of the introduction into Central Europe of the closely-relatedLyctus planicollis LeC. The article also points out that reintroduction of the European speciesLyctus linearis Gze., which was brought to North America a long time ago, could also occur.Practical morphological distinctions between two species (L. cavicollis andL. planicollis) as well as between the typical European speciesL. linearis and its morphacrassicollis Lesne as described from North America, are given.

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Mit 7 Tabellen und 4 Abbildungen     

Genetic diversity and adaptive traits of European versus American Douglas-fir seedlings

Due to its productivity and potential to adapt to the expected climate change, the Douglas-fir is one of the most important commercial non-native forest tree species in Europe. Currently, seeds from both non-native European and native American seed stands are used for plantations. In this study, we investigate European seed lots for their native origin (variety and potential geographic origin in America) and assess the adaptability, growth and survival potential of European versus American Douglas-fir seed lots. We compare the genetic diversity, morphological characteristics such as height (h), root collar diameter (rcd) and the ratio of h/rcd, and the timing of bud burst. We investigate 852 1-year-old seedlings from 10 different US and European seed lots representing 5 provenance regions which are sold in Germany and Austria. Seedlings are genotyped for 13 nuclear SSRs and analysed together with reference data set and standard genetic structuring and assignment methods. Adaptive traits of morphological characteristics and timing of bud burst of the seedlings are recorded and statistically analysed. The results show that the investigated European seedlings originate from recommended American native seed sources and represent both varieties and inter-varietal admixed individuals. European seedlings have a lower genetic diversity versus the American seedlings and native populations. They show significant differences in the adaptive traits such as morphological characteristics and timing of bud burst. According to the genetic diversity indices, certified North American Douglas-fir seed sources should be preferred for planting in Central Europe.     

Extent,distribution and origin of non-native forest tree species in Europe

ABSTRACT

The management of non-native tree species in European forests has a long history, but the information on the current number and geographic distribution of these species in European forests is incomplete and scattered across various datasets. This study aims to perform an inventory of the most frequent non-native tree species growing in European forests and analyse their current extent, geographic distribution and geographic origin. Our results show that at least 150 non-native tree species are currently growing in European forests and provenance trials. The genera represented by largest number of species are Eucalyptus, Pinus, Acacia and Abies. Species growing at the largest areas are Robinia pseudoacacia (2.44 million ha), Eucalyptus globulus (1.46 million ha), Picea sitchensis (1.16 million ha) and Pseudotsuga menziesii (0.83 million ha). In total, non-native tree species in Europe are found in an area of approximately 8.54 million ha, or 4.0% of the European forest area, and the five most abundant species account for up to 77% of this area. The largest number of these 150 species were introduced from North America (71), followed by Asia (45) and Australia (20). North American species occupy by far the largest area.     

Growth of Phlebiopsis gigantea in wood of seven conifer species

Heterobasidion parviporum and Heterobasidion annosum are widely distributed root‐rot fungi that infect conifers throughout Europe. Infection of conifer stumps by spores of these pathogens can be controlled by treating fresh stumps with a competing non‐pathogenic fungus, Phlebiopsis gigantea. In this study, growth of three Latvian strains of P. gigantea and the biological control agent ‘Rotstop’ strain was evaluated in stem pieces of Norway spruce, Scots pine, lodgepole pine, Douglas‐fir, Weymouth pine, Siberian larch and Sitka spruce. The growth rates of one H. parviporum and one H. annosum isolate were also measured in the same stem pieces. The growth rate of P. gigantea varied greatly in wood of different conifer species. It was higher in the three pine species, lower in Norway spruce and lowest in Sitka spruce and Siberian larch, and in Douglas‐fir, this fungus did not grow. The largest area of wood occupied by P. gigantea was in lodgepole pine. Growth of Latvian isolates of P. gigantea in the wood of Pinus and Picea species was comparable to that of the Rotstop isolate. Consequently, stump treatment with local P. gigantea isolates should be recommended. However, our results suggest that Douglas‐fir stump treatment against Heterobasidion by P. gigantea may be ineffective and other stump treatment methods should be considered.     

Influence of branch autonomy on fruit,scaffold, trunk and root growth during stage III of peach fruit development

We studied the influence of branch autonomy on the growth of reproductive and vegetative organs by establishing different patterns of fruit distribution within and between large branch units (scaffolds) in mature peach trees (Prunus persica (L.) Batsch cv. 'Elegant Lady'). Different patterns of fruit distribution were established by defruiting either whole scaffolds (uneven fruit distribution between scaffolds; US) or several selected hangers (small fruiting branches) per tree (uneven fruit distribution between hangers; UH). The effects of these patterns were compared with the effects of an even fruit distribution treatment (EVEN) in which fruits were thinned to achieve maximum uniformity of fruit distribution within the canopy. The desired fruit loads were obtained by differentially thinning the remaining bearing parts. On a tree basis, the response of mean fruit mass to fruit load was strongly affected by fruit distribution. The steepest mean fruit mass to fruit load relationship was found in US trees, whereas the relationship in UH trees was intermediate between the US and EVEN trees. On a scaffold basis, differences in fruit size between EVEN and US trees with similar fruit loads, though statistically significant, were relatively small, indicating that scaffolds were almost totally autonomous with respect to dry matter partitioning to fruit during the final stage of peach fruit growth. Hangers also appeared to exhibit significant autonomy with respect to the distribution of dry matter during the final phase of fruit growth. Branch autonomy was evident in scaffold growth: defruited scaffolds in the US treatment grew more than fruited scaffolds, and fruit distribution treatments had little impact on scaffold cross-sectional area on a tree basis. On the other hand, as observed for fruit growth, branch autonomy did not appear to be complete because the fruited scaffolds grew more in US trees than in EVEN trees under heavy cropping conditions. However, the effect of fruit distribution occurred only over short distances, and was negligible on organs located farther away from the source of heterogeneity (fruits), such as the trunk and roots.     

Evaluation of the biological control agent Rotstop in controlling the infection of spruce and pine stumps by Heterobasidion in Latvia

The biological control agent Rotstop^® composed of a suspension of spores of Phlebiopsis gigantea (Fr.) Jül. is widely used for protecting conifer stumps from aerial infection by Heterobasidion species. The efficacy of Rotstop application on Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) stumps was determined in several locations and at different seasons in Latvia. Mean efficacy in controlling natural infection by Heterobasidion spp. in spruce stumps was 64%, calculated on the basis of number of infected stumps, and 89%, calculated on the basis of area of infected wood on sample discs cut from the stumps. Corresponding proportions for pine were 82% and 95%. The results show that Rotstop can be successfully used for stump treatment in Latvia, although improved efficacy is desirable, particularly in spruce. A Latvian isolate of P. gigantea, selected from numerous isolates in preliminary tests, was included in one experiment and was shown to be as effective as the Rotstop isolate. In untreated spruce stumps Heterobasidion spp. and P. gigantea were present in the same stump three times more frequently than in untreated pine stumps. Heterobasidion spp. infection in untreated spruce stumps was low when P. gigantea covered more than 10% of stump dissection.     

Utility pole decay

Summary The Basidiomycetes associated with decay in pine, Douglas-fir, and cedar utility poles within various geographic regions of North America were investigated. On the basis of 313 isolations from these poles, 9 fungi appeared to be of major importance in internal pole decay in the United States. These fungi were the following: Lentinus lepideus, Lenzites saepiaria, L. trabea, Peniophora A., P. gigantea, and Poria radiculosa in pine, and L. lepideus, Poria carbonica, P. monticola, and P. xantha in Douglas-fir poles. Lentinus lepideus was overwhelmingly predominant in pine poles, whereas Poria carbonica was similarly predominant in Douglas-fir poles. Lenzites trabea was the fungus most often isolated from cedar poles; however, it is believed to be associated primarily with shell rot of cedar poles and to be of little significance, therefore, in causing internal decay of cedar. The fungi associated with western red-cedar and red and jack pine poles in Canada are listed, although the frequency of their occurrence is not included.This work was in cooperation with the Navy Department, Naval Facilities Engineering Command.The author is indebted to the following for furnishing cultures and pole sections or both for culturing or for information on the identities of the species of decay fungi found in Canadian poles: Joe Clark and John Kulp, U. S. Forest Products Laboratory, Madison, Wisconsin; Robert Graham and John Mothershead, Forest Research Laboratory, Oregon State University, Corvallis, Oregon; O. Floyd Hand, Bonneville Power Administration, Vancouver, Washington; John Shields, Canadian Forest Products Laboratory, Ottawa, Ontario; and John Roff, Canadian Forest Products Laboratory, Vancouver, British Columbia. The author is especially indebted to members of the Forest Disease Laboratory, Laurel, Maryland, for their invaluable aid in identifying representative cultures of many of the fungi discussed in this investigation.The Laboratory is maintained at Madison in cooperation with the University of Wisconsin.     

Variation in properties of Phlebiopsis gigantea related to biocontrol against infection by Heterobasidion spp. in Norway spruce stumps

The properties of 64 heterokaryotic strains of Phlebiopsis gigantea, isolated mostly from Norway spruce stumps, were tested for asexual spore production, growth rate and competitive ability against Heterobasidion spp. on agar medium, and for growth rate in spruce wood. Eighteen isolates were also tested for the efficacy of control against Heterobasidion spp. in stem pieces of spruce. The results revealed high variation in traits between different P. gigantea isolates. The efficacy of control against Heterobasidion spp. and the growth rate in spruce wood were closely related to each other (r = 0.727, p < 0.001). These preliminary tests indicate that the growth rate of a P. gigantea strain in spruce wood is the most important characteristic determining its efficiency in controlling the infection and spread of Heterobasidion spp. in spruce stumps.     

Capability of the European and North American race of Gremmeniella abietina to hydrolyse polygalacturonic acid in vitro

Polygalacturonase was found to be one of the first enzymes secreted by a pathogen during infection. The polygalacturonic-acid hydrolysing activity was compared between the North American and the European race of Gremmeniella abietina in vitro. Isolates were grown in pure pectin media from which the enzyme activity was analysed. Altogether, 29 isolates were tested in five experiments (experimental runs in a growing chamber). The data were analysed using variance-component models that included fixed-race effects and random-experiment, isolate, flask and measurement effects. The European race secreted more polygalacturonic-acid hydrolysing enzyme than the North American race and the mycelial dry weight produced was smaller for the European race. The differences between races were of the same order of magnitude as the variation between isolates within races; variance components relating to experimental errors were quite large. No correlation was found between the activity and mycelial dry-matter production within the races. Logarithmic transformation removed the apparent racial differences in the variability of the activity and mycelial dry weight. Results from the additionally tested A- and B-type of Finnish isolates indicated differences in dry-matter production.     

Phylogenetic and differentiation analysis of the trunk rot pathogen Fomitiporia tsugina in North America

Fomitiporia hartigii s.l. is an important trunk rot pathogen of conifers throughout the Northern Hemisphere. In North America, this pathogen primarily attacks Tsuga, but is also found on Abies, Picea and Pseudotsuga. Previous research showed that isolates of F. hartigii from North America represent a distinct phylogenetic species, known as F. tsugina. However, that conclusion is based on limited data. To better understand the phylogenetic relationships of F. tsugina in North America, a phylogenetic analysis was performed using three loci (internal transcribed spacer, nuclear large subunit and tef1) with 23 isolates originating from the northern United States, Canada and central Europe. North American isolates formed a monophyletic group with significant statistical support, confirming previous reports that F. tsugina represents a unique phylogenetic species, distinct from European F. hartigii s.s. Population subdivision between isolates of F. tsugina from eastern (Massachusetts, Michigan, New Hampshire, New York and Wisconsin) and western (British Columbia, Colorado, Oregon and Washington) North America was investigated. Tests of gene flow and genetic differentiation based on region of origin detected significant variation (F_ST = 0.761; K_ST = 0.625, p < 0.01), suggesting gene flow between the two populations may be limited. Neutrality tests revealed significant, negative departures from the standard neutral model, which could indicate that a purifying or stabilizing selection has maintained low levels of polymorphisms in the population, perhaps favouring an advantageous phenotype. Further studies are required to better understand the occurrence of F. tsugina on Abies and Picea in boreal forests outside the natural range of Tsuga.     

Comparison of five strains of Phlebiopsis gigantea and two Trichoderma formulations for treatment against natural Heterobasidion spore infections on Norway spruce stumps

In an experiment established on three Norway spruce sites in southern Sweden, the ability of five strains of Phlebiopsis gigantea, including the commercial strain Rotstop®, and two Trichoderma formulations to control natural Heterobasidion spp. spore infections was compared. At each site 160 trees were felled, and the resulting stumps were treated with spore suspensions of the seven fungal preparations. Twenty stumps at each site were left untreated as control stumps. When sampled 9 months after treatment, two of the P. gigantea strains (1984 and 1985) were the most effective at preventing infection by Heterobasidion spp. The other three P. gigantea strains were less effective, and two Trichoderma formulations did not significantly reduce Heterobasidion spp. infections.     

The susceptibility of Asian,European and North American <Emphasis Type="Italic">Fraxinus</Emphasis> species to the ash dieback pathogen <Emphasis Type="Italic">Hymenoscyphus fraxineus</Emphasis> reflects their phylogenetic history

In Europe, common ash (Fraxinus excelsior) is being decimated because of the invasive fungus Hymenoscyphus fraxineus. In its native range in Asia this ascomycete is considered a harmless leaf associate of F. mandshurica and F. chinensis subsp. rhynchophylla. Field observations from Europe suggest that there is species-specific variation in disease susceptibility among European and North American Fraxinus species, but a wider comparison at the genus level has been missing so far. We assessed disease symptoms and pathogen apothecium development in 17 Fraxinus species from Asia, Europe and North America exposed to high infection pressure in a Danish arboretum. We also tested their susceptibility to pathogen infection through controlled stem and leaf inoculations and subsequently assessed the level of pathogen DNA by a qPCR assay. The results suggested the presence of a phylogenetic signal in disease susceptibility where closely related Asian, European and North American species in section Fraxinus had relatively high levels of H. fraxineus DNA in the leaves and supported high production of apothecia. Leaves from some North American species also contained relatively high levels of H. fraxineus DNA, supported moderate production of apothecia and developed lesions—stating the need to avoid introduction of H. fraxineus to North America.