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.     

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.     

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.     

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.     

Optimizing the Management of a Butt-rotted Picea abies Stand Infected by Heterobasidion annosum from the Previous Rotation

There are no instructions on the management of stands of Norway spruce [ Picea abies (L.) Karst.] in which butt rot caused by Heterobasidion annosum coll. (Fr.) Bref. was not discovered until the time of first thinning. A stochastic simulation model describing the spread of butt rot in a stand of Norway spruce was used with a new submodel describing the butt rot of young trees. Non-linear stochastic optimization was used to determine the most profitable management schedule of a young Norway spruce stand with butt rot. If the initial level of butt rot was 5% or 10%, or the stand was thinned in summer with stump treatment, one thinning with a rotation of 55 yrs was optimal. Two winter thinnings with a rotation of 61 yrs was optimal in an initially healthy stand. Optimizations indicated that silvicultural measures that decrease the transfer of H. annosum to next generation at clear-cuttings could be economically fulfilled.     

Serpula himantioides,Heterobasidion annosum and Calocera viscosa as butt rot fungi in a Danish Douglas‐fir stand

A scattered wind fall of 50 trees in a 46‐year‐old stand of Douglas‐fir (Pseudotsuga menziesii) revealed Serpula himantioides, Heterobasidion annosum and Calocera viscosa as decay causing fungi. Sixty‐four percent of the stumps had visible rot on the stump surface. Mainly, a central brown rot was seen, caused by either S. himantioides or C. viscosa combined with eccentrically placed spots of incipient H. annosum rot. Subsequently, the appearance of fruitbodies disclosed that in total 76% of the wind thrown trees were associated with one or more of the three decay fungi. S. himantioides was present in a surprisingly large number of trees (54%), and the rot column extended up to 2 m into the stem. This investigation represents the first known incidence of S. himantioides and C. viscosa causing root and butt rot in living conifers in Denmark.     

Transfer of the P-type of Heterobasidion annosum from old-growth stumps of Picea abies to Picea abies and Larix×eurolepis

The infection of Picea abies and Larix x eurolepis by Heterobasidion annosum was studied in felled trees in Sweden. Thinnings were carried out in two stands of L. x eurolepis, 15 and 20 years old, and in a 25-year-old stand of P. abies, established on a site heavily infected by H. annosum. The 15- and 20-year-old L. x eurolepis stands exhibited the greatest incidence of butt rot with infections amounting to 38 and 57%, respectively, of removed trees. The incidence of butt rot in the P. abies stand was only 5%. Heterobasidion annosum was the main butt rot causing fungus. All isolates of H. annosum were of the P-intersterility group. Transfer of H. annosum from old-growth P. abies stumps to trees felled in the thinning of the next generation was demonstrated by somatic incompatibility tests. However, the majority of infections could not be connected with decay already established in the previous generation. In L. c eurolepis, rot columns were frequent, including also several genotypes in the same stem, and typically positioned in the inner sapwood. The few infections of P. abies were situated in the heartwood.     

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.     

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.     

Community of Aphyllophorales and Root Rot in Stumps of Picea abies on Clear-felled Forest Sites in Lithuania

The community of Aphyllophorales fungi in stumps of Picea abies (L.) Karst. and the occurrence of root rot caused by Heterobasidion annosum were investigated at 38 clear-felled sites in Lithuania. Fruit-bodies were recorded on 36.0% of the 3924 examined stumps. The most common were Bjerkandera adusta, Phlebiopsis gigantea, Fomitopsis pinicola, Gloeophyllum sepiarium, Stereum sanguinolentum, Trametes zonata and H. annosum. All species occurred in stumps both with and without established root rot, but B. adusta, F. pinicola, G. sepiarium, T. zonata and Trichaptum abietinum occurred less often in stumps containing root rot, while H. annosum was encountered more often. Greater numbers of fruit-bodies were found on stumps cut in summer than on those cut in winter. The fruiting became most abundant 3-4 yrs after the trees were felled. The abundance of nearly all Aphyllophorales correlated positively with the stump diameter, and on larger stumps, fruit-bodies of several different fungi were more common. The number of stumps containing H. annosum root rot in different sites varied from 9.8% to 68.8%, and was 27.6% on average. The incidence of root rot correlated neither with the age nor with the density of the felled stand, but it correlated negatively with the proportion of deciduous trees within a stand.     

Urea treatment reduced Heterobasidion annosum s.l. root rot in Picea abies after 15 years

Stump protection using chemical or biological agents is the main control measure against root and butt rot caused by Heterobasidion annosum s.l. in northern and temperate conifer forests. Long-term effects of urea treatment of stumps are poorly known and here we describe a 15-year study of urea treatment on the rot incidence on Norway spruce (Picea abies). We also tested the effect of urea protection on tree growth and on the resistance of stands against strong winds. Four treatments were made in two replications in two first-rotation P. abies stands in southern Sweden; after first-thinning stumps were (i) treated with urea 35% (w/v), (ii) artificially infected with H. annosum conidia, (iii) half urea treated and half artificially conidia infected, (iv) untreated, therefore naturally infected. After 15 years, the trees were sampled at 20 cm above ground using an increment borer and observed for presence of rot and, following incubation, presence of H. annosum conidia. Tree growth was calculated by measuring the diameter before and after the treatment. Urea treated plots showed the lowest incidence of rot (3%) as well as of H. annosum incidence (0%). Conidia treatment showed the highest incidence of rot (68%), its incidence was higher than that observed in natural infection treatment (43%), but did not differ from that of the 50% conidia treatment (47%). On about 30% of the rotted trees we observed conspicuous H. annosum fruiting. We did not observe growth reduction associated with tree rot. H. annosum was the only fungus observed associated with rotted trees which suggest that it was responsible for most of the rot observed in the investigated plots. Urea treated plots showed the lowest incidence of windthrown trees, and 59% of the windthrown tree incidence among the plots was explained by the incidence of H. annosum. Urea can be regarded as a reliable long-term protection method against root and butt rot of Norway spruce.     

Precommercial thinning of beech (Fagus sylvatica L.): Early effects of stump height on growth and natural pruning of potential crop trees

Abstract

This paper reports the early effects of stump height on the growth and natural pruning of potential crop trees after precommercial thinning of a young stand of naturally regenerated beech (Fagus sylvatica L.) in Denmark. The experiment comprises five treatments based on combinations of three grades of precommercial thinning and three stump heights. Treatments include the unthinned control, thinning only of whips and wolf trees (using low stumps), and thinning for potential crop trees using low, medium or high stumps. All treatments were replicated three times. Stump heights averaged 10, 90 or 230 cm, respectively. Pretreatment stem number (live trees) varied from 17,500 to 41,000 ha^?1. In the potential crop tree treatment, post-treatment stem number ranged from 4750 to 9500 ha^?1. Following two growth seasons, the quantity of stump regrowth increased with increasing stump height, the rate of stump regrowth increased with increasing stump height, the diameter growth of potential crop trees increased with decreasing stump height, the increase in stand height did not depend on stump height or post-treatment stem number, and the natural pruning of potential crop trees increased with increasing stump height. It remains to be seen whether these trends hold in the long run, and whether additional economic return from the increase in wood quality with increasing stump height compensates for the reduction in diameter growth.     

Heterobasidion annosum root rot in Picea abies: Modelling economic outcomes of stump treatment in Scandinavian coniferous forests

Abstract

The economic outcomes of stump treatment against spore infections of the root rot pathogen Heterobasiodion annosum s. l. were analysed based on simulations in four stands typical of Swedish forestry and forest management: (A) Norway spruce [Picea abies (L.) Karst.] stand on former agricultural soil (SI?=?32), (B) Norway spruce stand (SI?=?26) on forest land; (C) Mixed stand of Norway spruce and Scots pine (Pinus sylvestris L.) (SI?=?24) with only H. parviporum present, i.e. no infection of Scots pine and no possibility of interspecies spread of disease between hosts; and (D) same as C, but H. annosum s. str. (Fr.) Bref and H. parviporum Niemelä & Korhonen present, i.e. interspecies spread of disease possible. Models for disease development, growth and yield and cross-cutting were used in the simulations. The simulated decay frequency in Norway spruce trees ranged between 2 and 90%. Stump treatment at the previous final felling and in all thinning operations was profitable at interest rates 1 and 3% in stands A, B and D, but not in stand C. In stand C, no stump treatment at all or treatment in thinnings only gave the highest net future value. Implications for stump treatment in practical forestry are discussed.     

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.     

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.     

Relationship between stump treatment coverage using the biological control product PG Suspension,and control of Heterobasidion annosum on Corsican pine,Pinus nigra ssp. laricio

The relationship between the proportion of the stump surface covered by the biological stump treatment agent PG Suspension, containing Phlebiopsis gigantea and its efficacy against the pathogen Heterobasidion annosum sensu stricto was studied during a first thinning of Corsican pine (Pinus nigra ssp. laricio) in Thetford Forest, UK. PG Suspension was manually applied to 100%, 75%, 50% or 0% of the surface of 150 stumps. Spores of H. annosum were inoculated onto 75 of the stumps, and the remaining stumps exposed to natural airborne spore deposition. The relationship between coverage and efficacy was found to be quantitative. Covering all the stump surface with PG Suspension completely excluded the pathogen, whereas stumps not treated with PG Suspension (the 0% treatment) became infected with H. annosum. Partial (75%) PG Suspension coverage resulted in the pathogen colonizing 40% of stumps following artificial inoculation with H. annosum, and just 7% of stumps exposed to ambient H. annosum spore infection. Decreasing levels of coverage allowed increasing areas of the stump surface to be colonized by H. annosum. Some small gaps in coverage were closed by lateral growth of P. gigantea, but it is recommended that operators aim for full stump coverage to give complete protection against H. annosum.     

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.     

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.     

Modelling incidence of root rot in picea abies plantations in Southern Sweden

An empirical model is presented to forecast the incidence of root rot at stand level. In addition, the impact of different thinning programmes on the incidence of root rot is evaluated. The model is based on data from 152 permanent forest research plots of pure Picea abies in southern Sweden, within which the incidence of root rot at stump height in thinned trees has been recorded after each thinning since 1950. In total, about 20 000 stumps have been studied. According to simulations with the model, areas previously used as fields or for grazing are particularly susceptible to root rot, while old hardwood sites are less susceptible. Furthermore, simulations with the model imply that the earlier, the harder or more often a stand is thinned, the faster will be the development of root rot.     

Occurrence of Heterobasidion basidiocarps on cull pieces of Norway spruce left on cutting areas and in mature spruce stands

Fruiting of Heterobasidion on cull pieces and stumps of Norway spruce was investigated in cutting areas and mature spruce stands located in southern Finland. Cull pieces of variable size and showing butt rot were left on three clear‐cut areas and in one thinned stand. Additionally, a part of the cull pieces was transported to mature forest sites with closed canopy. During the succeeding 3–4 years the cull pieces were investigated annually for sporocarps of Heterobasidion, and the area of actively sporulating pore layer of each sporocarp was measured. Root bases of spruce stumps in the logging areas were excavated and sporocarps found on the stumps also measured. At the onset of the experiment, Heterobasidion spp. were isolated from 76% of the cull pieces showing butt rot; 85% of the isolates were identified as H. parviporum and 15% as H. annosum s.s. During the following 3–4 years sporocarps were found on 20% of the 1938 cull pieces where Heterobasidion butt rot was initially detected visually. Sporocarp formation was promoted by advancement of butt rot, increasing cull piece diameter and end‐to‐end ground contact, but restricted by the colonization of the cull piece by Stereum sanguinolentum. Between‐site differences were significant but could not be explained by differences in tree cover. At the end of the investigation period the average sporulating area of Heterobasidion sporocarp per metre of cull piece was higher than the average sporulating area per stump at three of four logging sites. Hence, leaving cull pieces containing Heterobasidion butt rot at logging areas in southern Finland can considerably increase local production of Heterobasidion spores.