The action of allicin against Armillaria spp. in vitro

The effect of allicin (a stabilized garlic extract product) at five different concentrations (0, 20, 30, 50 and 100 mg/l) was studied in vitro on the growth rate of 100 isolates of Armillaria gallica and A. mellea. Isolates were obtained from 41 host genera growing in gardens located in 39 counties in the United Kingdom. Agar plugs of the actively growing Armillaria isolates were added to the centre of malt agar plates infused with allicin, and radial mycelium growth was measured on days 7, 14 and 21. The total number of rhizomorphs and length of rhizomorphs were also measured. Relative growth rates were calculated as the growth rate relative to the controls (0 mg/ l). The relative growth of each isolate at each allicin concentration was used to estimate EC₅₀ values for A. mellea and A. gallica populations as well as individual isolates. EC₅₀ values for both Armillaria spp. increased over time. The mean EC₅₀ values for A. mellea of 16.0, 26.4 and 102 mg/l (days 7, 14 and 21, respectively) were higher than those for A. gallica (8.8, 7.9 and 11.0 mg/l) and probably relate to the more aggressive nature of A. mellea. Isolates with higher EC₅₀ values were also more likely to produce more rhizomorphs. At allicin concentrations of 20 and 30 mg/l, the production of rhizomorphs and the growth rates of A. mellea isolates were stimulated, when compared to the control treatments. From this study's findings, it appears that the field use potential of allicin is limited, due to better inhibition of the less virulent A. gallica, than the more aggressive A. mellea.     

Effects of woodland structure on breeding bird populations in stands of coppiced lime (Tilia cordata) in western England over a 10-year period

Bird populations were estimated in three adjacent stands ofsmall-leaved lime (Tilia cordata) at Shrawley Wood, Worcestershire,between 1984 and 1993. The stands were old coppice, recent coppicecut in the late 1970s and thinned coppice that had been singledprobably in the 1970s. It is thought that the stands were allpreviously coppiced in the 1940s. The study area presented anunusual opportunity to (1) assess effects of woodland structureon bird populations in the absence of any major differencesin tree species, (2) to compare the dynamics of bird populations,and hence assess habitat preferences, within large samples ofeach habitat. Overall densities of birds were consistently lowestin the old coppice and generally highest in the recent coppice.The singled coppice supported the least diverse bird communitiesas measured by a simple diversity index and the expected numbersof species within standardized samples of individuals. Warblers(Sylviidae) were strikingly more abundant in recent coppicethan other habitats, being virtually absent from the old coppice.Wrens were by far the most abundant species in the singled coppice.There was strong evidence that the old coppice was the leastpreferred habitat of wren and robin. The most striking changeduring the study was the collapse of the warbler populationsin the recent coppice after some 11 to 12 years of coppice growth.This was predictable from previous studies, but the timing ofthe warbler decline occurred later than expected and was probablya consequence of the relatively slow canopy closure of the coppiceat Shrawley Wood. Differences between stands, and changes overthe 10 years within stands, were interpreted in terms of habitatstructure and effects of winter weather on survival of residentspecies. The findings are discussed in relation to managementof abandoned coppice; it is suggested that for conservationpurposes thinning should be regarded as a complementary approach,alongside coppicing and non-intervention.     

Molecular‐based identification and phylogeny of Armillaria species from Serbia and Montenegro

Armillaria causes problems of root rot, kill trees and decay wood in the forests of Serbia and Montenegro, but the species involved have not hitherto been identified. The aim of this study was to identify field isolates collected on 25 localities. Identification was based on restriction fragment length polymorphism (RFLP) analysis of intergenic spacer 1 (IGS1) region and comparisons of IGS1 sequence with those available on NCBI database. Phylogenetic analysis was performed on sequence information from selected isolates to determine possible interrelationships between isolates with different banding patterns and previously identified tester isolates of five European Armillaria species. Five Armillaria species were identified in 90 isolates obtained from forests in Serbia and Montenegro. Armillaria gallica was most frequently isolated, followed by A. cepistipes, A. mellea, A. ostoyae and A. tabescens; two isolates remained unidentified. Restriction digestion of IGS1 amplification products with AluI produced 10 RFLP patterns. Patterns G4 (400, 250, 180) for A. gallica and pattern X (400, 180, 140) for isolates 74 and 79 are reported for the first time in European isolates. Eight RFLP patterns were observed after restriction with TaqI. Two patterns each were observed for A. ostoyae and A. gallica, and one each for A. cepistipes, A. mellea, A. tabescens and isolates 74 and 79. Parsimony analyses based on the IGS1 region placed the isolates into four clades: one including A. mellea, the second containing A. gallica–A. cepistipes isolates, while isolates of A. ostoyae and A. borealis were in the third clade. Armillaria tabescens differed from all annulate species. Phylogenetic analysis supported the conclusion that European Armillaria species are closely related and separated from a common ancestor in the near past. According to this survey five European Armillaria species are present in the forests of Serbia and Montenegro, while A. borealis is not present in the studied ecosystems.     

Identification and distribution of Armillaria species associated with an oak decline event in the Arkansas Ozarks

Forests in the Ozark Mountains of northern Arkansas recently experienced a widespread oak decline event. Armillaria, a root rot fungus, has been associated with other oak decline events and may have been an important contributing factor to tree mortality in this event. Although Armillaria has been identified from the Ozark Mountains in Missouri, it has never been investigated in the Arkansas Ozarks. Molecular diagnostic techniques were used in this study to identify species of Armillaria present on roots removed from dead trees of two common oak species, northern red oak, Quercus rubra L., and white oak, Q. alba L., from three geographic areas and on three topographic positions – ridges, south‐ and west‐facing benches. Armillaria(A. mellea, A. gallica or A. tabescens) was identified from 31% of root samples taken from 102 trees in seven of nine sample plots. Armillaria mellea, occurred most often (20 samples, both oak species on seven plots) followed by A. gallica (10 samples, northern red oak only on four plots), and A. tabescens occurred twice (on northern red oak in a single plot). Thus, all three Armillaria species occurred on northern red oaks while A. mellea was the only species recovered from white oaks. Results varied by topographic position with samples from tree roots on ridges having the fewest positive identifications, one of 29. West‐facing benches had the highest positive samples with 20 of 41 testing positive and trees on south‐facing benches were intermediate with 11 of 32 samples from infected trees. This study documents the occurrence of three species of Armillaria in the Arkansas Ozarks and their association with oak mortality resulting from an oak decline event coupled with a red oak borer, Enaphalodes rufulus, outbreak. Further, it documents some potential variation in host/pathogen combinations and forest site conditions.     

Penetration and colonization of oak roots by Armillaria mellea in Wisconsin

The level of Armillaria mellea activity on root systems of living and herbicide-killed oâks was assessed. Epiphytic rhizomorphs were almost always present on living and dead oak roots. After an oak was killed or completely defoliated, A. mellea usually penetrated directly through the root-collar bark and spread rapidly in the cambium. Sapwood in the root collar and main lateral roots was colonized within 2 years. The production of thizomorphs from this region was assessed on trees that had been killed by herbicide sprays 2 to 14 years before sampling.     

Geographical distribution of the Armillaria species in The Netherlands in relation to soil type and hosts

The geographical distribution of the annulate Armillaria species was studied in The Netherlands during the period 1983–1992. Armillaria gallica (incl. A. cepistipes), A. mellea and A. ostoyae appear to be widespread, the first two species being rather common on broad-leaved hosts growing on clay and loess soils, and the third species common on both broad-leaved and coniferous hosts on acid sandy soils. The distribution of the Armillaria species encountered was primarily determined by soil type. From a silvicultural point of view, A. ostoyae is the most important species, being pathogenic and occurring most frequently on sandy soils, the soil type predominantly used for forestry in The Netherlands.     

Phylogenetic analysis of Japanese Armillaria based on the intergenic spacer (IGS) sequences of their ribosomal DNA

To determine the phylogenetic positions of two new species, Armillaria jezoensis and Armillaria singula, and one new subspecies, Armillaria mellea suhsp. nipponica, the nucleotide sequences of the intergenic spacers (IGS) of their ribosomal DNA were investigated, and compared with those of tour other Armillaria species from Japan, and those of nine Armillaria species from Europe and North America. We conclude that Armillaria jezoensis, and Armillaria singula belong to the Armillaria gallica cluster as Armillaria cepistipes, Armillaria gallica and Armillaria sinapina from Japan. Two isolates of Armillaria ostoyae from Japan were placed within the Armillaria ostoyae cluster. Armillaria mellea subsp. nipponica had an IGS sequence as long as the IGS of Armillaria mellea from Europe and North America. However, the IGS sequences of Armillaria mellea subsp. nipponica, whose basidium base lacks a clamp connection could not be satisfactorily aligned with the IGS sequences of other species possessing this morphological feature.     

Identification of Armillaria species in Japan using PCR-RFLP analysis of rDNA intergenic spacer region and comparisons of Armillaria species in the world

Armillaria species from Japan were characterized using polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) of the intergenic spacer region-1 (IGS-1) of ribosomal DNA (rDNA). Eleven different digestion patterns by restriction endonuclease Alu I were found among 70 isolates of seven Armillaria species in Japan. Isolates within Armillaria nabsnona, A. ostoyae, A. cepistipes, and Japanese biological species E showed the same Alu I digestion patterns. Five Alu I patterns were detected for A. gallica, three patterns for A. mellea, and two patterns for A. tabescens. Seven Armillaria species in Japan were clearly distinguished by using the profiles obtained when PCR products were digested with Alu I, Msp I, and Hae III restriction enzymes. There was considerable variability of Alu I restriction sites within the IGS-1 between the isolates of five Armillaria species, A. gallica, A. nabsnona, A. cepistipes, A. mellea, and A. tabescens, in Japan and those of their European and North American counterparts.     

Ecology of Armillaria species in managed forests and plantations in Serbia

The distribution of Armillaria species was investigated in Serbian forest ecosystems, in relation to the main host species attacked, forest‐types, geography and altitude. In total, 388 isolates were identified from 36 host species in 47 sites. Armillaria gallica was the most commonly observed species with the widest distribution and with an altitudinal range of 70–1450 m, it was the dominating Armillaria species in lowland alluvial forests and in Quercus and Fagus forests at higher elevations. Armillaria mellea occurred in Quercus spp. – dominated forests in the north and central regions at 70–1050 m. Sixty‐eight per cent of the A. mellea isolates were collected from living hosts, most commonly in declining conifer plantations. Armillaria ostoyae was distributed in the cooler coniferous forest types and plantations in the Dinaric Alps in the south of Serbia, at 850–1820 m. Armillaria cepistipes was found in the eastern and southern hilly and mountainous regions of the country, at 600–1900 m. Most isolates were obtained from conifers and rhizomorphs in the soil around decaying stumps. Armillaria tabescens was found only on dead oak material in the northern and eastern regions of the country at altitudes lower than 600 m.     

The production of rhizomorphs by Armillaria mellea from stumps

The production of rhizomorphs by Armillaria mellea from stumps samples collected in East Anglia was studied: those of broad-leaved trees produced greater yields than those of pines and, in the absence of poisoning, over a much longer period after felling. The ability of A. mellea to form rhizomorphs is often reduced after prolonged growth in pine tissues. The interaction of A. mellea and other fungi is briefly considered. Some implications of these observations for forestry practice are discussed.     

The use of glyphosate for the management of secondary coppice regrowth in a Eucalyptus grandis × E. urophylla coppice stand in Zululand,South Africa

In South Africa, commercial eucalypt stands may be regenerated through the stepwise reduction of coppice shoots following felling. The development of secondary coppice regrowth following these reduction operations results in competition with the remaining stems, possibly resulting in a reduction in growth. Currently, secondary coppice regrowth is controlled manually, or through spraying with glyphosate at 0.6% when ca. 0.75?m in height. No research has been conducted to determine whether alternative rates and/or timing of application are possible or cost-effective. In 2006 a trial was implemented in Zululand, South Africa on a recently coppiced stand of Eucalyptus grandis × Eucalyptus urophylla so as to optimise rates of glyphosate application (0.6%, 1.2% and 1.8%) for the control of secondary coppice regrowth at various heights (0.5, 1.0 and 1.5?m). Tree and secondary coppice regrowth variates were measured annually until rotation-end (9 years, 3 months). The volume of herbicide used, the number of occasions each treatment was sprayed and associated costs were combined to provide an indication of the most cost-effective treatment. Although there were no significant differences in final coppice yield for the various rates of glyphosate and timing of application (as assessed by the size of the secondary coppice regrowth) tested, treatment efficacy in terms of treating secondary coppice regrowth increased with an increase in the rate of glypho- sate applied (0% < 0.6% < 1.2% < 1.8%), especially when treated at either 1.0 or 1.5?m in height. The use of glypho- sate, irrespective of rate and/or timing of application (as assessed by secondary coppice regrowth height), proved to be more cost-effective compared with manual control or the spraying of the secondary coppice regrowth at 0.75?m height with 0.6% glyphosate. The most cost-effective treatment was 1.2% glyphosate applied when the secondary coppice regrowth was 1.5?m. If reduced herbicide use is a major criterion within a company portfolio, then the secondary coppice regrowth can be manually removed when 1.5?m.     

Comparison of taper functions between two planted and coppiced eucalypt clonal hybrids, South Africa

In addition to regeneration through seed, certain eucalypts are able to regenerate via the production of coppice shoots following felling, which can then be selectively thinned over time and managed as a coppice stand for the commercial production of timber. Little information could be found if tree form differs between coppiced (where one or two stems had been left per stump) and planted eucalypts, or whether different volume taper models would need to be developed. To determine if this was necessary, rotation-end stem taper data was collected from an Eucalyptus grandis × E. urophylla and an E. grandis × E. camaldulensis trial to compare volume taper equations for planted versus coppiced commercially grown Eucalyptus clonal hybrids. For treatment comparisons, taper data were collected from the parent crop (1R), the replanted crop (2R), as well as from coppiced stands where either a single (Cop_Sngl) or double stem (Cop_Dbl) had been left per stump. Stem taper functions used for treatment comparisons indicated differences between clones, as well as between treatment, with the models based on single stems (1R, 2R or Cop_Sngl) being significantly different from those containing two stems (Cop_Dbl). Despite any non-significant model differences (for example between 1R, 2R and Cop_Sngl single stem crops) the percentage magnitude of any bias in utilizable volume differences when comparing the individual models (between a 6.31% over-prediction to a 3.8% under-prediction) still needs to be taken into account as these differences may have importance in terms of the volume and product prediction.     

Growth characteristics and allometry of <Emphasis Type="Italic">Robinia pseudoacacia</Emphasis> as a silvopastoral system component

Demand for goat (Capra hircus) meat in the southeastern USA is steadily increasing as a result of preferences exhibited by ethnic communities. Feeding systems that include fodder trees can be developed to take advantage of the natural preference of goats for browse. Data were collected for 2 years on a 5-year old stand of Robinia pseudoacacia L. to evaluate growth characteristics for goat browse and to develop allometric relationships in a randomized complete block design (intra-row spacing 0.5 or 1 m and coppice height 0.25 or 0.50 m) replicated six times. Allometric equations were derived from destructive harvests of 68 trees. Main branch size was not affected by intra-row spacing when trees were coppiced at 0.25 m; however, when coppiced at 0.5 m, trees spaced at 1 m had more and larger branches (P < 0.05) than trees spaced at 0.5 m. Intra-row spacing did not influence tree herbage biomass when trees were spaced at 0.5 m whereas trees coppiced at 0.25 m and spaced at 1 m produced less woody and herbaceous biomass than trees spaced at 0.5 m (P < 0.05). A strong relationship (P < 0.0001) was found between tree herbage biomass and the number of main branches greater than 0.01 m (r ² = 0.80). The results, based on growth characteristics and tree herbage biomass production, suggest that Robinia pseudoacacia L. would be an excellent candidate as a silvopastoral component in the southeastern USA.     

Effects of coppicing on the root and stump carbohydrate dynamics in birches

Whole birch stems were cut off in order to determine how coppicing affects root and stump starch, glucose, fructose and sucrose concentrations and their correlation with shoot regeneration capacity. The Betula pubescens Ehrh. and B. pendula Roth studied included intact trees, trees that had been coppiced 8 years earlier, trees coppiced at the beginning of the experimental season, and birches that had been coppiced twice, 8 years earlier and at the beginning of the experimental season. Carbohydrate accumulation differed between 8 years earlier coppiced and intact trees. Recent coppicing clearly decreased the starch and sugar concentrations of the roots, which were often highest in the thin roots. The concentrations of these compounds in the stumps were always low, although the carbohydrate concentrations of stumps, in particular, correlated with shoot regeneration capacity. Starch was the most labile of the carbohydrates measured and most clearly reacted to coppicing. Differences in starch- and sugar-reserve dynamics indicate a difference between these birch species in the use and replenishing of root and stump reserves. This information may also be of help when the effects of other stresses, for example, severe animal damage or burning, on the regrowth of young birch stands are estimated.     

Distribution of Armillaria species in upland Ozark Mountain forests with respect to site,overstory species composition and oak decline

Armillaria root disease is a contributing factor to oak decline in the Ozark Mountains of central USA. We have identified Armillaria gallica, Armillaria mellea, and Armillaria tabescens in Quercus‐Carya‐Pinus forests of the region. Presence/absence patterns of each Armillaria species as well as all possible Armillaria species combinations were analysed by contingency tables and/or stepwise logistic multiple regressions with principal characteristics of the studied sites and forest stands, both quantitative and qualitative: geographic land‐type association, bedrock type, landform position, slope direction (aspect), soil type and soil surface stone cover, down woody debris, abundance and basal area of woody vegetation and decline mortality by species. Most decline mortality consisted of two red oak species (section Erythrobalanus, Quercus coccinea and Quercus velutina), which also were most sensitive to Armillaria infection. Site characteristics related to the distributions of Armillaria species and decline mortality were also related to the preponderance of Q. coccinea and Q. velutina, regional vegetation history (i.e. conversion of Pinus echinata stands to hardwoods), and the different strategies of territory acquisition and spread of the Armillaria species involved. The presence of A. gallica may reduce the activity of more virulent Armillaria species.     

The effect of coppice management on stump volume recovery in mechanized operations

Key message

Coppice stands result in slightly higher stump waste compared with planted stands, when felled mechanically by a harvester.

Context

The large demand for wood fibre requires efficient production and cost-effective practices throughout the supply chain.

Aims

The purpose of the study was to quantify the amount of volume lost to excessive stump height in coppiced and planted stands.

Methods

Stump height was measured on similar eucalypt stands that differed only for their origin: coppiced or planted. The study sample comprised of 543 planted stems and 851 coppice stems; of which 365 grew as double stems and 486 as single.

Results

Stump waste was highest for coppiced double stumps, smallest for coppiced single stumps and intermediate for planted tree stumps. All differences were statistically significant, but the difference between coppiced single stumps and planted tree stumps was much smaller (20%) than the difference between coppiced double stumps and the rest (220–260%). Regression analysis showed that stump waste volume increased with tree volume, and this effect was twice as large for coppiced double stumps compared with the other treatments. Stump waste seemed very small in both relative and absolute terms and is unlikely to offset the large benefits accrued through coppice management and mechanization.

Conclusion

Comparison with previous stump height studies indicates that the results obtained in this experiment for planted eucalypt may have general value and could be extended to other coppice stands, although with caution.

     

A comparison of the cost-effectiveness of different eucalypt cut-stump control management options to reduce competition from coppice regrowth during stand establishment in Mpumalanga,South Africa

Selected cut-stump (Cut_surface) and basal frill (Basal_frill) treatments were tested on Eucalyptus macarthurii cut stumps that had previously been coppiced multiple times. Treatments included the application of herbicide (triclopyr as an amine salt, 360 g L^?1), either to the cut-surface or to a basal frill (cuts made through the bark and into the cambium at the base of the tree). These treatments were integrated with non-chemical control methods, where seedlings were planted either between each stump row or within the row, to determine whether such treatments could further reduce the level of coppicing. Stump mortality for both the Basal_frill and Cut_surface treatments were equally effective, with 87% of the stumps killed when assessed at 14 weeks following initial treatment. No significant treatment differences were detected for the various growth variates of the stand at rotation end (stocking, basal area and volume). This can be attributed to a lack of competition from any coppice regrowth due to good control on each occasion. As there were no significant differences between the treatments, the Cut_surface treatment would be preferable due to the lower overall costs. However, for this method to be most effective, it is important that the herbicide be applied to the cut-surface of the felled tree within one day of felling. This may pose a safety issue, especially with mechanised felling of a stand. Further research on extending the period for application to the cut-surface could be beneficial. However, if reduced herbicide use is a prerequisite, coppice regrowth can be manually removed.     

Development of species‐specific PCR primers on rDNA for the identification of European Armillaria species

Attempts to design species‐specific PCR primers from six European Armillaria species in the ribosomal RNA genes are reported. Primers were developed on the basis of the nucleotide sequence variability of the internal transcribed spacers (ITS) and the intergenic spacer (IGS1) of the ribosomal DNA. Four sets of primers gave specific PCR products for Armillaria tabescens, Armillaria mellea and Armillaria ostoyae. However, due to the high sequence similarities between Armillaria borealis and Armillaria ostoyae and between Armillaria cepistipes and Armillaria gallica no species specific amplification was obtained for these taxa.     

Distribution and ecology of Armillaria species in Greece

Five Armillaria species were identified in a nationwide survey in Greece. Armillaria mellea was present in coniferous and broad-leaved forests in most of the areas examined, except the high altitudes (above 1100 m) of the mountains of north Greece. It was found to cause significant damage in fir forests as well as in fruit orchards and vineyards. Armillaria gallica was common in coniferous and broad-leaved forests in the high altitudes of central and northern Greece, predominating in the beech forests. The fungus was a weak parasite or a saprophyte of forest trees and was occasionally found on cultivated plants. Armillaria ostoyae was not found in southern and central parts of the country, but it has a wide distribution in the mountain forests of northern Greece and causes significant damage on fir, black pine, Scots pine and spruce. Armillaria cepistipes was recorded at high altitudes (1400–1800 m) on two mountains of northern Greece, mostly as a saprophyte in coniferous and broad-leaved forests. Armillaria tabescens was rare in the forests of Greece; it was found to cause disease in almond tree orchards.     

Stump infection by Armillaria in first-rotation conifers

Information about the entry of Armillaria into first-rotation pine and spruce stands was obtained by searching for infected stumps, rhizomorph systems or trees that had been killed. In pines Armillaria foci were very rare. In pure Norway spruce Armillaria lutea and A. mellea were detected in stumps but rhizomorphs did not extend into the soil; in Norway spruce mixed with oak, by contrast, A. lutea sometimes produced extensive rhizomorph systems. In Sitka spruce small groups of trees had been killed by A. ostoyae. All foci investigated in conifers contained different genotypes of Armillaria and probably originated from spore infection of stumps created by thinning. Some implications of these findings are discussed.