Summary A large database of invasive forest pathogens (IFPs) was developed to investigate the patterns and determinants of invasion in Europe. Detailed taxonomic and biological information on the invasive species was combined with country‐specific data on land use, climate, and the time since invasion to identify the determinants of invasiveness, and to differentiate the class of environments which share territorial and climate features associated with a susceptibility to invasion. IFPs increased exponentially in the last four decades. Until 1919, IFPs already present moved across Europe. Then, new IFPs were introduced mainly from North America, and recently from Asia. Hybrid pathogens also appeared. Countries with a wider range of environments, higher human impact or international trade hosted more IFPs. Rainfall influenced the diffusion rates. Environmental conditions of the new and original ranges and systematic and ecological attributes affected invasiveness. Further spread of established IFPs is expected in countries that have experienced commercial isolation in the recent past. Densely populated countries with high environmental diversity may be the weakest links in attempts to prevent new arrivals. Tight coordination of actions against new arrivals is needed. Eradication seems impossible, and prevention seems the only reliable measure, although this will be difficult in the face of global mobility.

Wounding of Norway spruce by inoculation with sterile agar, or agar containing the pathogenic fungus Ceratocystis polonica, induced traumatic resin duct formation in the stem. Visible anatomical responses occurred in the cambium 6-9 d post-inoculation. Near the inoculation site cellular proliferation, polyphenolic accumulation, and lignification were induced as a wound reaction to seal the damaged area. Five centimetres from the inoculation site cells in the cambial zone swelled and divided to form clusters. By 18 d post-inoculation, these cells began to differentiate into resin duct epithelial cells surrounding incipient schizogenous lumens. Mature axial traumatic ducts appeared by 36 d as a row of ducts in the xylem centripetal to the cambium. The ducts formed an interconnected network continuous with radial resin ducts. Parenchyma cells surrounding the ducts accumulated polyphenols that disappeared as the cells differentiated into tracheids. These polyphenols appeared to contain fewer sugar residues compared to those accumulating in the secondary phloem, as indicated by the periodic acid-Schiff's staining. The epithelial cells did not accumulate polyphenols but contained immunologically detectable phenylalanine ammonia lyase (EC 4.3.1.5), indicating synthesis of phenolics as a possible resin component. These findings may represent a defense mechanism in Norway spruce against the pathogenic fungus Ceratocystis polonica.

Dothistroma needle blight (DNB) is one of the most important diseases of pine. Although its notoriety stems from Southern Hemisphere epidemics in Pinus radiata plantations, the disease has increased in prevalence and severity in areas of the Northern Hemisphere, including Europe, during the last two decades. This increase has largely been attributed to expanded planting of susceptible hosts, anthropogenic dispersal of the causative pathogens and changes in climate conducive to disease development. The last comprehensive review of DNB was published in 2004, with updates on geographic distribution and host species in 2009. Importantly, the recognition that two species, Dothistroma septosporum and D. pini, cause DNB emerged only relatively recently in 2004. These two species are morphologically very similar, and DNA-based techniques are needed to distinguish between them. Consequently, many records of host species affected or geographic location of DNB prior to 2004 are inconclusive or even misleading. The objectives of this review were (i) to provide a new database in which detailed records of DNB from 62 countries are collated; (ii) to chart the current global distribution of D. septosporum and D. pini; (iii) to list all known host species and to consider their susceptibility globally; (iv) to collate Drenkhan et al. 410 |

Summary• Parasitism and saprotrophic wood decay are two fungal strategies fundamental for succession and nutrient cycling in forest ecosystems. An opportunity to assess the trade-off between these strategies is provided by the forest pathogen and wood decayer Heterobasidion annosum sensu lato.• We report the annotated genome sequence and transcript profiling, as well as the quantitative trait loci mapping, of one member of the species complex: H. irregulare. Quantitative trait loci critical for pathogenicity, and rich in transposable elements, orphan and secreted genes, were identified.• A wide range of cellulose-degrading enzymes are expressed during wood decay. By contrast, pathogenic interaction between H. irregulare and pine engages fewer carbohydrate-active enzymes, but involves an increase in pectinolytic enzymes, transcription modules for oxidative stress and secondary metabolite production.• Our results show a trade-off in terms of constrained carbohydrate decomposition and membrane transport capacity during interaction with living hosts. Our findings establish that saprotrophic wood decay and necrotrophic parasitism involve two distinct, yet overlapping, processes.

Dieback of European ash (Fraxinus excelsior), caused by the ascomycete Hymenoscyphus pseudoalbidus (anamorph Chalara fraxinea), started around 1992 in Poland and has since then spread over large geographical areas. By November 2010, the disease had been recorded in 22 European countries. The gradual expansion and high intensity of the ash dieback epidemic in Europe may suggest that H. pseudoalbidus is an invasive alien organism. In Norway, ash dieback was first reported in spring 2008, and a survey in early summer of the same year revealed that the disease had spread over large parts of the southern and eastern regions of the country. The distance from the southernmost to the northernmost infected stands was, at that time, about 400 km. Some old necrotic lesions were also observed, indicating that the ash dieback pathogen is likely to have been present in Norway since at least 2006. In 2009, a spore sampler was installed in a diseased ash stand at Ås, South‐Eastern Norway. Sampling started in late July and continued until late September. Large numbers of ascospores resembling those of H. pseudoalbidus were observed, with the maximum number of spores occurring from the end of July to mid‐August. The deposition of ascospores occurred mainly between 6 and 8 a.m. Ascospores are most likely to be the primary source initiating host infections and responsible for the rapid recent spread of H. pseudoalbidus in Europe.

Isolations of microorganisms were made from the sapwood of Norway spruce (Picea abies) (L.) Karst. infested by the spruce bark beetle Ips typographus L. The most important group of fungi isolated, the species of Ophiostomataceae, is treated. Ten species were isolated, of which Ophiostoma polonicum Siem. appears to be the most active invader of the sapwood. Three new species are described, Ophiostoma ainoae sp. nov., O. flexuosum sp. nov. and O. cucullatum sp. nov. and three new combinations are proposed, O. davidsonii (Olchow. & Reid.) comb. nov., O. europhioides (Wright & Cain) comb. nov. and O. sagmatospora (Wright & Cain) comb. nov.

The pathogenicity of two isolates of each of four bark beetle-associated blue-stain fungi was evaluated after mass inoculation of about 40-year-old Norway spruce trees (Picea abies). Trees were inoculated with a different isolate of each fungus in 1995 and 1996 at a density of 400 inoculations per m(2) in a 1.2-m-wide band on the lower bole (about 270 inoculations per tree). Trees were felled 15 weeks after inoculation. In 1995, Ceratocystis polonica was the only fungus that had stained the sapwood (56.3% of cross-sectional sapwood area). It induced five times longer phloem necroses, 21 times more dead cambium, and 11 times more dead phloem than any other fungus. In 1996, C. polonica induced less extensive host symptoms and an unidentified Ambrosiella sp. induced comparable symptoms to C. polonica in the phloem and cambium. No trees showed any foliar symptoms 15 weeks after inoculation, but six out of eight trees inoculated with C. polonica in 1995 had only 0 to 25% functional sapwood and probably would have died if felling had been delayed. This study confirms that C. polonica, an associate of the aggressive bark beetle Ips typographus, is pathogenic to Norway spruce. The pathogenicity of the Ambrosiella sp., which is associated with a nonaggressive bark beetle, seems moderate and varies between isolates. The two remaining fungi included in this study (Ophiostoma piceae and a dark fungus with sterile mycelium), which are associated with nonaggressive bark beetles, were nonpathogenic in both experiments. These results are consistent with the hypothesis that aggressive bark beetle species vector virulent fungi that may help them kill trees, but the results also show that some nonaggressive bark beetles may vector phytopathogenic fungi.

Fungi associated with five bark beetle species colonizing Norway spruce (Piceaabies (L.) Karst.) were isolated from beetle-inoculated logs. Ipstypographus L., an aggressive tree-killing bark beetle, was associated with a different range of blue-stain fungi than the nonaggressive Pityogeneschalcographus L., Polygraphuspoligraphus L., and Hylurgopspalliatus Gyll. The flora of the nonaggressive Ipsduplicatus Sahib. was similar to that of I. typographus. The pathogenic fungus Ceratocystispolonica (Siem.) C. Moreau, and other blue-stain fungi, were isolated in high frequency from inoculations with both Ips species. Pathogenic blue-stain fungi were absent, or isolated in low frequencies, from inoculations with the other nonaggressive beetle species. With the exception of I. duplicatus, these results support the hypothesis that aggressive bark beetles carry more pathogenic blue-stain fungi than other bark beetles and indicate that pathogenic fungi are important for aggressive bark beetles to kill trees.