Soil properties and tree growth on rehabilitated forest landings in the interior cedar hemlock biogeoclimatic zone: British Columbia

We studied operational landing rehabilitation programs in three forest districts of interior British Columbia (BC). Winged subsoiling and grass/legume seeding, followed by planting of lodgepole pine (Pinus contorta var. Latifolia) generally resulted in successful re-establishment of forest cover on landings. In the Boundary district, fifth year tree heights on landings were not significantly different from those in adjacent plantations, and fifth year growth increments were also similar despite evidence of delayed seedling establishment caused by cattle grazing damage. Trees on landings in the Kalum district were shorter than those in plantations, but fifth year increments were similar. Landings in the Kalum with >20% clay had lower stocking densities, tree heights, and probe depths compared to landings with <20% clay content. In the Kispiox district, average fifth year tree heights and fifth year increments were lower on landings than those on plantations. In the Kispiox district, landings with >20% clay content had shorter trees growing on them when compared to tree heights on landings with <20% clay. Landings in the Kispiox had the least probe depth of the three districts, and the greatest difference in height and increments between landings and plantations, supporting field reports of poor decompaction effectiveness there. In all districts, there was no forest floor present on landings 7 years after subsoiling, and cover of non-coniferous vegetation was lower than for plantations. Higher soil temperatures and lower soil moisture contents were recorded on landings relative to plantations in all districts during the growing season of 1998. Landing soils also had lower total C, N and mineralizable-N (min-N) than plantation soils in the Boundary and Kispiox, but such differences were not statistically significant in the Kalum. Reduced levels of total C, N and min-N on landings did not appear to have affected current foliar nutrient status of trees growing on these sites, as most nutrients appeared in concentrations considered adequate or only slightly deficient, except for foliar S on all sites. Average concentrations of N, P, K and S in foliage were either similar to or significantly higher for trees on landings relative to plantations, but these differences were attributed to dilution in plantation foliage rather than deficiency. Our results support the conclusion that operationally feasible techniques for soil rehabilitation can create conditions suitable for establishment of a new forest on sites that otherwise would be considered non-productive. Stocking levels and tree growth rates observed in our study are consistent with the conclusion that a commercial forest may be produced on some of the rehabilitated areas.     

Retention of canopy lichens after partial-cut harvesting in wet-belt interior cedar–hemlock forests,British Columbia,Canada

Old-growth cedar–hemlock forests of the interior wet-belt of British Columbia are rich in abundance and diversity of canopy lichens, but are subject to forest harvesting. If these distinctive canopy lichen communities are to be conserved, it is important to learn how they are affected by forestry practices, including partial-cutting techniques designed to retain old-growth attributes. The retention of canopy lichens after 30 and 70% partial-cut harvesting (immediate post-harvest and 2-year post-harvest measurements) was examined using direct canopy access methods (tree climbing) in two old-growth interior cedar–hemlock forest stands of the upper Fraser River valley. Mean lichen loading was generally lower in all treatment areas, including control sites, 2 years after harvesting. However, three of four lichen-sampling groups (cyanolichen, foliose, and Bryoria group lichens) did not show significant treatment effects (total lichen loading) by harvesting type (30, 70 or 100% retention), when data sets were standardized against initial post-harvest lichen loading. Only in Alectoria group lichens were treatment effects observed. Although cyanolichen loading was not significantly different in retained trees in most harvest blocks, many thalli, especially those near south-facing edges, were discolored, suggesting that future cyanolichen loading may be lower within partial-cut harvest blocks. Litterfall in general was greater during 2001/2002 than 2002/2003, even in the control units, suggesting weather-related differences between years. For Alectoria, there appeared to be a treatment-related pulse of litterfall that was more pronounced in the 70% removal than the 30% removal treatment unit. Deposition patterns of litterfall components in relation to edges of openings indicated a heightened level of Alectoria litterfall along edges. Treatment-related differences were absent or weak in other groups, and cyanolichen litterfall did not differ between years in any study area/treatment unit combination. These findings suggest that lichen retention in the residual stand of both partial-cut treatments was adequate to meet management goals, but also point to the necessity of future monitoring, as other edge effects (e.g. microclimate changes) influence future lichen growth and mortality.     

Snag longevity of Douglas-fir,western hemlock,and western redcedar from permanent sample plots in coastal British Columbia

Snags are important both as structural components and as animal habitat in forests, but abundance is often low and their dynamics poorly understood in young, managed stands. Using a large data set of 19,622 snags from permanent plots in second-growth forests of coastal British Columbia, we modeled snag longevity (time from tree mortality to snag fall) for three species: Douglas-fir (Pseudotsuga menziesii), western hemlock (Tsuga heterophylla), and western redcedar (Thuja plicata). Snag longevity was strongly related to species and snag size (diameter): the median snag longevity was 16 years for Douglas-fir, 11 years for hemlock and 5 years for redcedar. Western redcedar was predominantly in the subcanopy and its rapid fall rate was related to the small size of its snags. In addition to diameter, other attributes (height to diameter ratio, height, and live crown ratio before death) contributed significantly to models for one or two of the species. However, site level variables did not contribute significantly to any of the models. Snags greater than 50 cm diameter, especially Douglas-fir snags, have the potential for persistence well beyond 20 years in these second-growth forests, and could be important for wildlife.     

Factors limiting early growth of western redcedar, western hemlock and Sitka spruce seedlings on ericaceous-dominated clearcut sites in coastal British Columbia

A 3 year field and pot study was conducted to determine the effects of several biotic and abiotic factors on the early growth of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western redcedar (Thuja plicata, Donn) and Sitka spruce (Picea sitchensis, Bong. Carr.) seedlings established on 2- and 8-year-old clearcut sites previously occupied by old-growth western hemlock and western redcedar forests (referred to as younger and older CH, respectively), and on adjacent 2-year-old clearcut sites previously occupied by second-growth western hemlock and amabilis fir (Amabilis amabilis (Dougl.) Forbes) forests (referred to as younger HA) in coastal British Columbia. The objective of the study was to determine which factors are associated with the poor growth characteristic of the salal (Gaultheria shallon, Pursh) dominated CH clearcut sites.

No soil moisture deficits were measured on any of the three types of clearcut sites at any time of year. The best seeding growth was on the younger HA sites followed by the younger CH sites and then the older CH sites. The better growth on the younger HA sites was associated with a higher availability of N and P in the first 20 cm depth of the forest floor. No differences in matric soil water potential and pH, and only small differences in soil temperature were measured between the three types of clearcut sites. Complete removal of the competing vegetation on the younger and older CH sites resulted in an increase in conifer seedling growth and in the availability of N (22–40%) and P (15–32%); however, it did not affect cellulose decomposition and matric soil water potential, and increased soil temperature only slightly. Both western hemlock and Sitka spruce seedlings were very responsive to differences in nutrient availability measured between types of clearcut sites and planting treatments. In contrast, western redcedar was not responsive. All three conifer species had very high mycorrhizal colonization on the younger CH sites, and this was not altered by the removal of the competing vegetation (mainly salal).

These results suggest that the nutritional stress and poor growth of conifers on salal-dominated CH clearcut sites in coastal British Columbia can be explained by:(1) inherently low forest floor nutrient availability; (2) competition between salal and conifer seedlings for scarce nutrients and nutrient immobilization in salal; (3) declining nutrient availability a few years after clearcutting and slashburning.     

Epidemiology of Armillaria root disease in Douglas‐fir plantations in the cedar‐hemlock zone of the southern interior of British Columbia

Results are presented from several studies on the epidemiology of Armillaria ostoyae in Douglas‐fir plantations in the interior cedar‐hemlock (ICH) biogeoclimatic zone of British Columbia. Two plantations were monitored for mortality by A. ostoyae and other agents for 35 years after establishment. In these and other plantations ranging in age from 7 to 32 years, one or more of the following factors were determined: source of inoculum, mode of spread and characteristics of lesions on roots of excavated trees; symptom expression in relation to tree age and damage to the root system and years from initial infection to death on trees killed by the fungus. Mortality from A. ostoyae began in both plantations about 5 years after planting, reaching 30% in one and 11% in the other after 35 years. The spatial pattern of mortality was similar to that reported from New Zealand, France and South Africa; however, the temporal pattern differed, beginning later and, instead of declining, continuing at a nearly constant rate to the present. To age 10, nearly all infections were initiated by rhizomorphs; as plantations aged, the proportion of infections occurring at root–root contacts increased. In seven plantations, in moist and wet subzones of the ICH, from 23 to 52% of Douglas‐firs had root lesions, with the higher incidences occurring on moist sites. The occurrence of aboveground symptoms, reduced leader growth and basal resinosis, was related to the percentage of root length colonized by A. ostoyae on trees with more than 30% of root length killed. Average time from infection to death increased from 1 to 2 years at age 6 to 22 years at age 33. The outlook for timber yield from Douglas‐fir plantations in the cedar‐hemlock zone is discussed. Management alternatives for reducing damage from A. ostoyae when regenerating sites are reviewed.     

Sheep vegetation management for controlling competing vegetation in young conifer plantations in the central interior of British Columbia,Canada

In British Columbia, sheep vegetation management (SVM) is a relatively new technique; thus, limited data are available for examining its benefits on conifer growth. We collected field data from young mixed-conifer plantations of lodgepole pine (Pinus contorta) and hybrid spruce (Picea glauca × P. engelmannii) in blocks which had been grazed in 2005 and 2006 as well as from ungrazed blocks. Our main objective was to determine if sheep grazing had a significant effect on the growth of hybrid spruce. We measured stem diameter at 15 cm height above the root collar (D15) and cumulative internodal length (IL). Cumulative IL was calculated from the node corresponding to the year 2002–2010 (positions 2–10). The height to diameter ratio (height to the base of the leader (position 10)/D15; HDR) was calculated to determine if seedlings prioritized growth in terms of height or diameter. There was no significant grazing treatment effect on D15 and HDR when compared to the control treatment; however IL was significantly affected after the second grazing treatment in 2006 (position 6) and became increasingly important with time (positions 7–10). Based on our results, we suggest that SVM could be an effective method for controlling competing vegetation and thus, increase annual tree growth for areas with similar site characteristics as those found within the plantations sampled.     

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.     

Climate and site characteristics affect juvenile trembling aspen development in conifer plantations across southern British Columbia

In southern British Columbia, juvenile trembling aspen is managed primarily as a competitor with conifers rather than for its ecological and economic value. As a result, brushing treatments have been applied on a widespread basis and this practice is likely to continue in the near future. Given the potential for climate change to affect our valuation of aspen, we require a better understanding of factors that affect its development, its competitive ability with conifers and its responses to brushing. We used data from 11 aspen management experiments to examine the influence of climate and site factors on aspen height, cover, and density in 17–24 year-old control stands and 9–16 years after manual cutting or girdling. Models explained 64% and 89% of the variation in aspen height in control and manually brushed stands, respectively, but were poor for girdling. Increasing length of the frost-free period was associated with increasing aspen height in control stands, whereas drier summer conditions on cool aspects favoured height growth of aspen suckers following manual cutting. We also examined the influence of climate and site factors on three simple competition indices that describe the height and density of aspen relative to conifer height, and then tested how well these indices predicted conifer growth. The density of aspen taller than conifers accounted for 39% of the variation in lodgepole pine diameter and the ratio of aspen/conifer height accounted for 33% of the variation in Douglas-fir height, suggesting that aspen competition was only moderately important to conifer growth. Our findings imply that aspen may become more productive with warmer summers provided it is not limited by summer moisture availability and that mixed stand management is a viable option in southern interior stands.     

The interaction between competition in interior Douglas-fir plantations and disease caused by Armillaria ostoyae in British Columbia

Interior Douglas-fir trees in plantations were assessed for size differences related to the level of diseased neighbours infected with Armillaria ostoyae. The four Douglas-fir stands studied ranged from 25- to 34-year-old, and represented the oldest accessible planted stands in the Interior Cedar Hemlock (ICH) ecosystem in British Columbia. Twenty-three to 25, 10-m radius plots were established in each stand. The spatial coordinates, total height, and diameter at breast height of all live and dead trees in the plots were recorded. Subject trees whose competitors were contained in the 10-m radius plots were also identified. Trees were pulled out of the soil using a mechanical excavator and the root systems were surveyed for evidence of infection by A. ostoyae. Stem disks were taken from each tree at 1.3 m above the ground for a determination of basal area. Increasing proportion of diseased trees in the plots resulted in less total plot basal area, but did not affect the mean basal area or height. Individual subject tree basal area was negatively related to the level of disease in surrounding competitors, opposite to expectations; however, diseased subject trees had reduced height and basal area compared to disease-free subject trees. Increasing competition reduced both the height and basal area of the trees, while regular distribution of all trees increased both total and mean plot basal area but not height. Disease incidence at the plot level and in individual subject trees was mainly affected by the neighbourhood conditions in which it grew, and was also related to disease intensity in the tree root systems. Although disease may alter resource partitioning among trees, the utilization of these resources is mostly limited by the increasing disease incidence as the stands age, the higher probability of larger trees being diseased with time, the occurrence of dead trees in clumps, and the high probability that dead trees will eventually infect live neighbours. The widespread belowground incidence of A. ostoyae in the ICH, its rapid colonization of stumps, and its wide host range can reduce site potential in managed stands.     

Comparison of carbon-stock changes,eddycovariance carbon fluxes and model estimates in coastal Douglas-fir stands in British Columbia

Background: The global network of eddy-covariance(EC) flux-towers has improved the understanding of the terrestrial carbon(C) cycle, however, the network has a relatively limited spatial extent compared to forest inventory data and plots. Developing methods to use inventory-based and EC flux measurements together with modeling approaches is necessary evaluate forest C dynamics across broad spatial extents.Methods: Changes in C stock change(ΔC) were computed based on repeated measurements of forest inventory plots and compared with separate measurements of cumulative net ecosystem productivity(ΣNEP) over four years(2003 – 2006) for Douglas-fir(Pseudotsuga menziesii var menziesii) dominated regeneration(HDF00), juvenile(HDF88and HDF90) and near-rotation(DF49) aged stands(6, 18, 20, 57 years old in 2006, respectively) in coastal British Columbia. ΔC was determined from forest inventory plot data alone, and in a hybrid approach using inventory data along with litter fall data and published decay equations to determine the change in detrital pools. These ΔC-based estimates were then compared with ΣNEP measured at an eddy-covariance flux-tower(EC-flux) and modelled by the Carbon Budget Model- Canadian Forest Sector(CBM-CFS3) using historic forest inventory and forest disturbance data.Footprint analysis was used with remote sensing, soils and topography data to evaluate how well the inventory plots represented the range of stand conditions within the area of the flux-tower footprint and to spatially scale the plot data to the area of the EC-flux and model based estimates.Results: The closest convergence among methods was for the juvenile stands while the largest divergences were for the regenerating clearcut, followed by the near-rotation stand. At the regenerating clearcut, footprint weighting of CBM-CFS3 ΣNEP increased convergence with EC flux ΣNEP, but not for ΔC. While spatial scaling and footprint weighting did not increase convergence for ΔC, they did provide confidence that the sample plots represented site conditions as measured by the EC tower.Conclusions: Methods to use inventory and EC flux measurements together with modeling approaches are necessary to understand forest C dynamics across broad spatial extents. Each approach has advantages and limitations that need to be considered for investigations at varying spatial and temporal scales.     

Comparison of carbon-stock changes,eddycovariance carbon fluxes and model estimates in coastal Douglas-fir stands in British Columbia

Background: The global network of eddy-covariance(EC) flux-towers has improved the understanding of the terrestrial carbon(C) cycle, however, the network has a relatively limited spatial extent compared to forest inventory data and plots. Developing methods to use inventory-based and EC flux measurements together with modeling approaches is necessary evaluate forest C dynamics across broad spatial extents.Methods: Changes in C stock change(ΔC) were computed based on repeated measurements of forest inventory plots and compared with separate measurements of cumulative net ecosystem productivity(ΣNEP) over four years(2003 – 2006) for Douglas-fir(Pseudotsuga menziesii var menziesii) dominated regeneration(HDF00), juvenile(HDF88and HDF90) and near-rotation(DF49) aged stands(6, 18, 20, 57 years old in 2006, respectively) in coastal British Columbia. ΔC was determined from forest inventory plot data alone, and in a hybrid approach using inventory data along with litter fall data and published decay equations to determine the change in detrital pools. These ΔC-based estimates were then compared with ΣNEP measured at an eddy-covariance flux-tower(EC-flux) and modelled by the Carbon Budget Model- Canadian Forest Sector(CBM-CFS3) using historic forest inventory and forest disturbance data.Footprint analysis was used with remote sensing, soils and topography data to evaluate how well the inventory plots represented the range of stand conditions within the area of the flux-tower footprint and to spatially scale the plot data to the area of the EC-flux and model based estimates.Results: The closest convergence among methods was for the juvenile stands while the largest divergences were for the regenerating clearcut, followed by the near-rotation stand. At the regenerating clearcut, footprint weighting of CBM-CFS3 ΣNEP increased convergence with EC flux ΣNEP, but not for ΔC. While spatial scaling and footprint weighting did not increase convergence for ΔC, they did provide confidence that the sample plots represented site conditions as measured by the EC tower.Conclusions: Methods to use inventory and EC flux measurements together with modeling approaches are necessary to understand forest C dynamics across broad spatial extents. Each approach has advantages and limitations that need to be considered for investigations at varying spatial and temporal scales.     

Factors affecting a sheep vegetation management system in British Columbia, Canada

A silvopastoral agroforestry system involving sheep is an important silvicultural issue. We examined the factors that affect the continuation or discontinuation of sheep vegetation management (SVM) in British Columbia. A questionnaire was used to conduct interviews with a randomly selected sample of contractors and farmers who were involved in SVM in British Columbia (BC). Based on a stepwise discriminant analysis, we found that the profitability of SVM, and the practitioners' commitment (number of ewes owned by the farmer or contractor), experience (number of years the farmer or contractor has practiced SVM on public lands in BC) and trust or confidence (number of years the farmer or contractor has consistently worked for the landlord) contributed to the farmer's or contractor's continuation or discontinuation of SVM practice. The factors we defined seemed to be validated (corroborated) by a complex of determinants voiced by various experts concerning the relationship between SVM and the forestry economy in BC, the scale of sheep farming in BC, the relative absence of an agricultural infrastructure to support sheep farming in BC, and the continuity of SVM from year to year. Further investigation into these factors and others is required to advance our understanding of SVM in British Columbia. This revised version was published online in June 2006 with corrections to the Cover Date.     

Recovery of N-urea 10 years after application to a Douglas-fir pole stand in coastal British Columbia

The long-term fate of fertilizer N in forest ecosystems is poorly understood even though such information is critical for designing better forest fertilization practices. We studied the distribution and recovery of ¹⁵N (4.934 atom% excess)-labelled fertilizer (applied as urea at 200 kg N ha⁻¹) 10 years after application to a 38–39-year-old Douglas-fir (Pseudotsuga menzeisii (Mirb.) Franco) stand in coastal British Columbia. The urea was applied in the spring (May 1982) or fall (November 1982). Sampling was conducted in October 1992, and we found that after 10 years, there were few differences between the fall and spring fertilizer applications in total N and ¹⁵N distribution within the tree and forest ecosystem. On average total fertilizer-N recovery was 59.4%; about 14.5% of the applied-N was recovered in the trees including coarse roots, with foliage containing 41% of the labelled-N recovered in the aboveground tree biomass. Tissue ¹⁵N remained mobile and could be transferred to new growth. Soil recovery was 39.8%, which had decreased from 57.0% at a previous 1-year sampling, with an average loss of 3.0% per year from the mineral soil and 3.7% from the litter layers. However, it appears that there was little continuing tree uptake. While short-term effects of fall vs. spring urea application were previously reported, there were no long-term effects on either stand productivity or fertilizer use efficiency, suggesting that if fertilization is properly done, timing of fertilization is not a critical issue in terms of maximizing fertilizer use efficiency for the coastal Douglas-fir forest we studied. Our results also highlight the high capacity of this ecosystem to retain externally applied inorganic N over the long-term, the importance of maximizing nitrogen uptake in the first year, and also of the continuing need to develop new approaches to overcome the generally low efficiency of forest N fertilization.     

Mapping the environmental limitations to growth of coastal Douglas-fir stands on Vancouver Island, British Columbia

Coastal Douglas-fir (Pseudotsuga menziesii spp. menziesii (Mirb.) Franco) occurs over a wide range of environmental conditions on Vancouver Island, British Columbia. Although ecological zones have been drawn, no formal spatial analysis of environmental limitations on tree growth has been carried out. Such an exercise is desirable to identify areas that may warrant intensive management and to evaluate the impacts of predicted climate change this century. We applied a physiologically based forest growth model, 3-PG (Physiological Principles Predicting Growth), to interpret and map current limitations to Douglas-fir growth across Vancouver Island at 100-m cell resolution. We first calibrated the model to reproduce the regional productivity estimates reported in yield table growth curves. Further analyses indicated that slope exposure is important; southwest slopes of 30 degrees receive 40% more incident radiation than similarly inclined northeast slopes. When combined with other environmental differences associated with aspect, the model predicted 60% more growth on southwest exposures than on northeast exposures. The model simulations support field observations that drought is rare in the wetter zones, but common on the eastern side of Vancouver Island at lower elevations and on more exposed slopes. We illustrate the current limitations on growth caused by suboptimal temperature, high vapor pressure deficits and other factors. The modeling approach complements ecological classifications and offers the potential to identify the most favorable sites for management of other native tree species under current and future climatic conditions.     

Effects of root zone trenching on soil nitrogen dynamics in Japanese cedar and cypress plantations

The effects of root exclusion and planted tree species on soil nitrogen (N) dynamics were examined at two plantations, one planted with Japanese cedar and the other with Japanese cypress. We set up ten 1 × 1 × 0.2-m-deep trenched sites and ten untrenched control sites at each plantation. We measured the pool size and leaching of inorganic N at each site for 2 years and the net N mineralization 1 and 2 years after trenching. Despite similar soil conditions, the cedar plantation showed higher net N mineralization than the cypress plantation. Stopped tree uptake of N was expected to cause an increased pool size and leaching of inorganic N at the trenched sites. Nevertheless, we found no significant increase in those variables at both plantations. The trenched cypress sites showed no decrease in the net N mineralization during the 2 years after trenching. However, the net nitrification at the trenched cypress sites increased remarkably at the deeper horizons in comparison with that at the control sites. Enhanced nitrification might result from improved ammonium availability through root exclusion. Net N mineralization at the trenched cedar sites decreased more than 60% compared with that at the control sites 2 years after trenching. Higher nitrification potential at the cedar plantation and enhanced nitrification potential at the trenched cypress sites never resulted in increased leaching of N, due to added fine root litter which acted as an immobilization agent for excess N, thus preventing N loss.     

Pathogenicity and virulence of Armillaria sinapina and host response to infection in Douglas‐fir,western hemlock and western redcedar in the southern Interior of British Columbia

The frequency of infection, lesion characteristics and anatomical changes in phloem and cambial tissues caused by Armillaria sinapina were studied on inoculated trees of Douglas‐fir, western hemlock and western redcedar and compared with results of A. ostoyae inoculations previously reported on the same host species. Similar percentages of inoculations resulting in infection of roots on the three hosts indicate that A. sinapina and A. ostoyae are equally pathogenic. Armillaria ostoyae was more virulent than A. sinapina as demonstrated by fungal exudates from A. ostoyae inoculum blocks, which appeared to cause lesions on roots; the higher frequency at which lignified impervious tissue (IT) and necrophylactic periderm (NP) developed in bark and following cambial invasion, compartmentalization; the large proportion of roots that showed no visible host response; the large zones of IT formed under continuous stimulation by A. ostoyae advancing in inner bark; and the high frequency of breaching of NP barriers. Spread of A. sinapina mycelium in host species appeared slower than that of A. ostoyae, particularly in Douglas‐fir and western hemlock. In western redcedar, A. ostoyae induced stronger host responses than those following invasion by A. sinapina, which included further expansion of the induced rhytidome response, traumatic phloem resin duct formation and higher numbers of polyphenolic parenchyma comprising its barrier zone. Where damage by A. sinapina ensued, it was always associated with high inoculum potential. The ecology of virulent and less virulent species of Armillaria in natural forests is discussed.