Effect of planting Scots pine around Norway spruce stumps on the spread of Heterobasidion coll.

Management of a Norway spruce stand planted on a site infected by Heterobasidion coll. is problematic because the fungus spreads vegetatively from the colonized stumps of the previous generation to the new seedlings. Growing of mixed stand with more resistant tree species has been suggested to decrease the economic losses caused by butt rot in Norway spruce trees. The mechanistic simulation model Rotstand describing the spread of Heterobasidion coll. in coniferous stands of southern Finland was used to study the effect of planting Scots pines around colonized clear‐felling stumps of Norway spruce of the previous generation. Planting of Scots pines in clusters around colonized stumps markedly decreased the butt rot of Norway spruce trees at the age of 20 years and at clear felling. If the same number of Scots pines were planted randomly, the effect was weak. When the average diameter of colonized clear‐felling stumps was 30 cm, a Scots pine circle with a radius of 3 m resulted in the highest soil expectation value (SEV) at 2% discounting rate, whereas with 40‐cm stump diameter, a 4‐m radius produced the highest SEV. When the proportion of Heterobasidion parviporum in the old colonized stumps was 50% instead of 95%, planting pines around colonized stumps still clearly decreased the butt rot at the age of 20 years and in final felling.     

Presence of Heterobasidion infections in Norway spruce stumps 6 years after treatment with Phlebiopsis gigantea

Natural colonization by the root and butt rot causing fungi Heterobasidion spp. on Norway spruce (Picea abies) stumps following thinning and treatment with the biological control agent Phlebiopsis gigantea was investigated on three sites in southern Sweden 6 years after treatment. The fully treated stumps and control stumps were excavated and sampled to compare the survival of Heterobasidion spp. in the long term. Six years post‐treatment, 47 and 11% of untreated and treated stumps, respectively, had Heterobasidion infection. There was no difference in the relative infected area in discs collected from the butt and the roots for the different treatments. Control efficacy was 83% for treated stumps. After 6 years, there were no apparent differences between the remaining infections in treated compared with those in untreated stumps regarding the number of colonies, their size or relative infection area. Although infections, 3 months after treatment with P. gigantea, were significantly fewer and smaller than in untreated stumps, Heterobasidion inoculum can survive for at least 6 years in the stump and, when it does, constitute a risk for neighbouring trees.     

Spatial distribution and persistence of Heterobasidion parviporum genets on a Norway spruce site

Spatial distribution of Heterobasidion genets over a period of ca 50 years in two successive generations of Norway spruce (Picea abies) was unravelled. The genets were first identified in 1993 in a naturally regenerated 43‐year‐old spruce stand that had been thinned the previous winter. Heterobasidion parviporum was found in 17.5% of the old stumps of the previous spruce generation. Nine genets were identified on the study plot; seven of them were present in old stumps of the previous tree generation and two only in the new spruce generation. Eighteen spruce trees of the new generation were infected, 15 of them by vegetative growth of genets originating from the old stumps. The study plot was investigated again in 2005. No new genets had been established after thinning, and three old genets had died out. The remaining genets had infected five new trees, most likely from the thinning stumps of diseased trees. At the age of 56 years, 16.1% of the residual spruces were infected by Heterobasidion. The results of this study suggest that if spore infection to stumps of spruce can be prevented, the decay frequency caused by H. parviporum will not necessarily increase in successive generations.     

Effectiveness of treatment of Norway spruce stumps with Phlebiopsis gigantea at different rates of coverage for the control of Heterobasidion

The natural establishment of the root and butt rot causing fungus Heterobasidion annosum s.l. on Norway spruce (Picea abies) thinning stumps treated with Phlebiopsis gigantea was investigated on seven sites in southern Sweden. The trees were cut during summertime and the stumps were treated with different patterns simulating the effect of mechanical stump treatment with a single‐grip harvester. Sampling was conducted 3 and 12 months after treatment. At both samplings, the best control was obtained when 100% of the stump surface was covered by P. gigantea: in contrast, untreated control stumps showed the highest incidences of H. annosum s.l. infection at both sampling times. However, 30 and 26% of the fully covered stumps at the first and second samplings, respectively, were diseased, and question the efficacy of treating Norway spruce stumps with this biological control agent in Sweden.     

Optimizing the management of a Picea abies stand under risk of butt rot

A simulation model was developed to predict the growth of a Norway spruce stand under risk of butt rot caused by Heterobasidion annosum stump infection and logging injuries. The simulation model was distance‐dependent; tree growth was predicted with a distance‐dependent model, and the spread of butt rot through root contacts depended on tree location. Infection of stumps and injured trees, and the spread of butt rot in the stand were stochastic processes whereas tree growth and mortality were treated as deterministic processes. The simulation model was used with the nonlinear optimization algorithm of ooke and J eeves (J. Assoc. Comput. Mach, 8, 212–229, 1961) to find the most profitable management schedule for an even‐aged, young stand. Optimization used four different stump infection rates and two spreading capacities from infected stumps. The profitability was evaluated by the expected soil expectation value (SEV) at a 3% interest rate. Two thinnings, both in winter‐time, and hence without H. annosum infections, resulted in the highest SEV. If any stump infection by H. annosum occurred, only one thinning and a shortened rotation were suggested. The optimal thinning rate tended to decrease but also large trees were removed with the increasing infection rate. With one thinning during a rotation, stump treatment was profitable above a stump infection rate of 10%     

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.     

Swiss stone pine trees and spruce stumps represent an important habitat for Heterobasidion spp. in subalpine forests

In the Western Italian Alps (WIA), the three European species of the forest pathogen Heterobasidion spp. can coexist in the same area. Heterobasidion parviporum Niemelä & Korhonen and Heterobasidion abietinum Niemelä & Korhonen are normally found in areas with a significant presence of their respective primary hosts, spruce (Picea spp.) and fir (Abies spp.). The host/niche occupied by Heterobasidion annosum (Fr.) Bref. in the region still remains unclear. Although Scots pine (Pinus sylvestris), a major host for this fungal species in other parts of Europe, is abundant in the region, little or no evidence of disease caused by H. annosum is visible in this tree species. Two different, but not mutually exclusive, hypotheses can explain the presence of H. annosum: (1) Scots pines are infected but largely asymptomatic and (2) H. annosum has adapted to different hosts. An analysis of Heterobasidion species was performed in two natural, mixed‐conifer forests using traditional isolation techniques and novel direct molecular diagnosis from wood. In a subalpine stand of mixed spruce (Picea abies), larch (Larix spp.), and Swiss stone pine (Pinus cembra), 18 naturally infected spruces and larches only yielded H. parviporum. A Swiss stone pine in the same stand was extensively colonized by both H. parviporum and H. annosum. In a second subalpine stand, an analysis of 18 spruce stumps and nine Swiss stone pine stumps yielded both H. parviporum and H. annosum isolates. Pine stumps had been mostly colonized by H. parviporum prior to tree felling, suggesting that this species may be secondarily infected by the locally predominant Heterobasidion species (i.e. H. parviporum). Results of our analysis also indicated that primary colonization of spruce stumps (e.g. through basidiospores) was caused by both H. parviporum and H. annosum, while secondary infection of such stumps was mostly because of H. parviporum.     

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.     

Efficacy of different concentrations of Rotstop® and Rotstop®S and imperfect coverage of Rotstop®S against Heterobasidion spp. spore infections on Norway spruce stumps

Two Rotstop^® preparations, one containing a Phlebiopsis gigantea strain from Finland (Rotstop F) and the other one from Sweden (Rotstop S), were used in different concentrations to treat thinning stumps of Picea abies against spore infections by Heterobasidion spp. in southern Sweden. Trees were cut on three sites during the summer of 2004, and 285 stumps were treated manually ensuring 100% coverage. Spore concentrations in the treatment suspensions were ca 5 × 10⁶ and 10 × 10⁶ spores/l, and approximately 10 ml of suspension was applied per 100 cm² of stump surface. An additional 31 stumps on one of the sites were treated mechanically with Rotstop S; in this treatment the spore concentration was high, about 20 × 10⁶ spores/l, but the coverage was incomplete. Three months later there was a significant reduction in frequency and relative areas of Heterobasidion spp. infections on stumps with manual treatment compared with the untreated stumps. However, there was no significant difference between the preparations or the concentration of active ingredient in terms of their reduction of Heterobasidion infection. Mechanical treatment with incomplete coverage failed to control infection. Therefore, in conditions of moderate Heterobasidion spore load in the environment there seems to be no reason to use higher concentrations of P. gigantea in the treatment of spruce stumps.     

Heterobasidion annosum: Development of Butt Rot following Thinning in two young First Rotation Stands of Norway Spruce

Severe butt rot caused by Heterobasidion annosum (Fomes annosus)developed in two stands of first rotation Norway spruce on oldagricultural land within 11 years of first thinning. At SiteI (Devon), 11 per cent of thinning stumps and 23 per cent ofclear felled stumps were found to be decayed when examined soonafter the crop was felled at age 37 years. It is estimated thatbetween 4.4 per cent and 6.6 per cent of potential volume waslost on clear felling. Significant butt rot had also developedin a 33-year-old stand of Norway spruce in Site II (Aberdeenshire)11 years after a Scots pine nurse crop was removed. Here, thedisease was found to be widespread on an area of 2 ha, and 73per cent of trees had butt rot. Decay extended 3 m up the stems,and occupied significant areas of the butt. In both cases, thepresence of butt rot in the crops led to premature clear felling,which resulted in additional loss of potential revenue. It seems most likely that the disease became established followinginfection of thinning stumps by airborne spores of the fungus:at Site I, the thinning stumps were not protected whilst atSite II it would appear that a chemical or biological treatmentapplied to the pine stumps failed. The subsequent rapid developmentof the disease was due, in part, to a variety of environmentaland edaphic factors.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

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.     

Incidence of Root and Butt Rot in Consecutive Rotations of Picea abies

The incidence of butt rot in two consecutive rotations of Norway spruce [Picea abies (L.) Karst.] in 28 permanent sample plots at four different sites in Denmark was evaluated. Incidence of butt rot was estimated by visual examination of stumps at final felling of the previous rotation and by examination of bore cores taken at the butt from a random sample of trees before first thinning of the subsequent rotation. There was no correlation between the incidence of butt rot at final felling of the previous rotation of Norway spruce and the incidence of butt rot at first thinning of the subsequent rotation of Norway spruce. The incidence of butt rot at final felling was between 19 and 100%, and at first thinning between 0 and 20%. The S-form of Heterobasidion annosum (Fr.) Bref. was the most commonly found decay-causing organism at all sites. Root systems of 28 trees without decay at stump height in the present rotation were excavated to estimate the incidence of root rot. Heterobasidion annosum was found in only one root. Resinicium bicolor (Alb. & Schw. ex Fr.) Parm. was found in 25% of the excavated root systems. The result of the study shows that the incidence of butt rot at first thinning of Norway spruce is not necessarily higher on sites where the previous rotation was heavily infected than on sites where infection in the previous rotation was low.     

Predicting the activity of Heterobasidion parviporum on Norway spruce in warming climate from its respiration rate at different temperatures

We studied the effect of climate warming on Heterobasidion root rot in boreal forests by measuring respiration activity of pure cultures of Heterobasidion parviporum in Norway spruce (Picea abies) sawdust and by linking these data to temperature data obtained from three spruce forests located along a north‐south transect stretching from northern Germany to northern Finland. The pure cultures applied in this investigation were homokaryotic, but in a separate investigation, we found no significant difference between the activity of homo‐ and heterokaryotic isolates. We also found that the temperature response curves of growth and respiration rates of this fungus were similar and propose that respiration reflects the general activity of H. parviporum. The respiration data were scaled up to annual cumulative respiration activity using daily temperature measurements from soil and air in the spruce forest sites. The annual respiration activity of H. parviporum showed a linear relationship with the average annual air temperature. An increase in the annual air temperature by 5°C would raise the annual activity of H. parviporum in spruce roots in northern Finland, southern Finland and northern Germany by 91%, 53% and 40%, respectively. This increase remains below the predicted increase in forest growth in northern Finland but exceeds considerably the predictions for southern Finland. According to the previous literature, a number of other decay fungi show a similar activity response to temperature as H. parviporum, suggesting that this result can be generalized to decay fungi with similar ecological habits.     

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.     

Controlling and predicting the spread of heterobasidion annosum from infected stumps and trees of picea abies

Two Norway spruce stands with heavy infections of Heterobasidion annosum were clear‐cut in 1957 and 1959 in Sweden. The stumps were extracted, the soil sifted to remove most of the roots, and young Norway spruce were planted. After 25 and 28 years, H. annosum had infected 1 % and 2% of trees on plots where stumps had been removed and 17 % and 12% of the trees on control plots, respectively. Several of the H. annosum clones fruiting on old‐growth stumps were also detected in decayed, standing trees. The same fungal clone was found to be infecting adjacent trees from several old‐growth stumps. In addition to old stumps, stumps from recent thinnings and diseased living trees were traced as infection sources. Their relative importance in spreading disease was estimated. Disease risk predictions based on the distance of a tree from various infection sources correlated well with observed frequencies of rot.     

Hosts and distribution of the intersterility groups of Heterobasidion annosum in the highlands of Greece

The intersterility groups of 127 pure cultures of H, annosum collected from different host trees in 12 mountain areas in Greece were identified. The F group commonly caused butt rot on the fir species Abies cephalonica and Abies borisii regis. It was the only type of H. annosum found in pure fir forests. The P group caused serious root rot in pinus sylvestris stands in north-eastern Greece. In more southern mountain areas it often colonized stumps of pinus nigra but seldom killed this tree species. The S group was found in natural forests of Picea abies in northern Greece, causing butt rot of spruce. In mixed forests, the intersterility groups of H. annosum were found relatively often in stumps of tree species other than their main hosts, although some host preference seemed to occur also in stump colonization.     

Root and butt rot of Todo fir (Abies sachalinensis) caused by Heterobasidion annosum s.l. in Hokkaido,Japan

The occurrence and symptoms of root and butt rot were examined in a 35 × 30 m plot of 68‐year‐old Todo fir plantation in Hokkaido, Japan. Forty‐seven percent of the cut stumps were decayed and 52% of the decayed stumps showed similar decay characteristics with yellowish orange to light brown colouration and expanded pockets in the heartwood. Morphological characteristics of the pure cultures isolated from the decay were similar to the cultures isolated from basidiocarps of Heterobasidion annosum sensu lato, found on fallen logs outside of the research site. Also DNA analysis based on the combined data set of three gene loci (glyceraldehyde 3‐phosphate dehydrogenase, heat shock protein 80–1 and elongation factor 1‐alpha genes) showed that the isolates from the decay are included in the same clade with the Japanese H. annosum s.l. isolates. They form a subclade to H. parviporum (the European S group of H. annosum s.l.). This is the first report of molecular determination of H. annosum s.l. isolated from root and butt rot in a plantation in Japan.     

New records of Heterobasidion parviporum in China

Twenty‐one basidiocarp specimens of Heterobasidion annosum s.l. were collected in six forest areas in western and central China, from host trees Picea schrenkiana, Abies fabri, Abies fargesii and Larix griffithii. Single‐spore cultures were isolated from the basidiocarps and identified by mating tests. All the specimens proved to belong to the species Heterobasidion parviporum. The results show that this fungus has a wide distribution in China. It seems to be a less aggressive pathogen in China than in Europe, although reliable data on the occurrence of Heterobasidion root rot in coniferous forests of China are so far lacking.     

Incidence of Heterobasidion annosum in precommercial thinning stumps in coastal British Columbia

Coniferous stumps in 83 stands in coastal British Columbia were sampled 3-5 years after precommercial thinning. The percentage of stumps and surface area colonized by Heterobasidion annosum were determined for 25 stumps of each species in each 5-cm diameter class present in each stand. There were significant differences among species in the percentages of stumps and surface area colonized, with Douglas-fir (Pseudotsuga menziesii) having the lowest values, amabilis fir (Abies amabilis) and Sitka spruce (Picea sitchensis) the highest and western hemlock (Tsuga heterophylla) being intermediate. For stumps of each species 5–20 cm in diameter, both the percentage of stumps and surface area colonized increased with increasing diameter. In stumps that were grafted to an adjacent tree, there was decreased incidence of H. annosum for Douglas-fir and Sitka spruce and increased incidence for western hemlock and amabilis fir. There were trends in the percentage of stumps and area colonized for season of thinning and biogeoclimatic subzones with the values for most species decreasing as the amount of precipitation increased. Colonization of precommercial thinning stumps by H. annosum occurs throughout the coastal region of British Columbia, and this will increase the amount of inoculum and will likely increase the incidence of butt rot. The results of this study suggest that the increase in inoculum can be minimized by thinning before age 15, by cutting only trees less than 10 cm in diameter and by thinning during low risk seasons.