Susceptibility of red fir and white fir to fir dwarf mistletoe (Arceuthobium abietinum) in California

Fir dwarf mistletoe (Arceuthobium abietinum, Viscaceae) is a damaging parasite of true firs (Abies spp.) in California. In the Sierra Nevada Mountains, fir dwarf mistletoe consists of two special forms (formae speciales, f. sp.) which are morphologically similar, but exhibit a high degree of host specificity: A. abietinum f. sp. concoloris whose principal host is Sierra white fir (Abies lowiana) and A. abietinum f. sp. magnificae which only parasitizes red fir (Abies magnifica). Although field observations and cross‐inoculation studies have consistently supported the extreme host specialization demonstrated by these special forms of fir dwarf mistletoe, little quantitative data from mixed red fir/white fir stands infested with fir dwarf mistletoe have been available. Therefore, I sampled 20 mixed conifer stands dominated by red and white fir and infested with f. sp. concoloris (8 stands) or f. sp. magnificae (12 stands) from near Mount Shasta to the southern Sierra Nevada Mountains near Sherman Pass. Infection of red and white fir was evaluated using temporary circular plots (0.012 ha) established around large, severely infected trees and with rectangular transects placed within fir dwarf mistletoe‐infested stands. In each plot and transect, live trees >5 cm in diameter at breast height were assigned a dwarf mistletoe rating (6‐class system). A total of 55 plots and 51 transects were established in the 20 stands, and over 1,900 red firs and 1,700 white firs were sampled. Ninety‐nine per cent of the red firs and 98% of the white firs were infected in stands infested with f. sp. magnificae or f. sp. concoloris, respectively. In contrast, no infection of red fir was observed in stands infested with f. sp. concoloris and no infected white firs were observed in stands infested with f. sp. magnificae. These data provide further support that the special forms of fir dwarf mistletoe are extremely host specific in California.     

The influence of prescribed crown fire on lodgepole pine dwarf mistletoe (Arceuthobium americanum) populations 33 years post‐fire

Dwarf mistletoes (Arceuthobium spp.) are a group of obligate, hemiparasitic plants that infect numerous species in the Pinaceae in North America. Wildland fire is considered to be the primary natural agent influencing the population and distribution of dwarf mistletoes across landscapes. Based on this understanding, prescribed fire has been suggested as a potential method for dwarf mistletoe sanitation and control; however, experimental work has primarily focused on prescribed surface fire. In this study, we report long‐term impacts of three experimental crown fires on dwarf mistletoe severity in infested lodgepole pine stands in Colorado 33 years post‐fire. The three fires achieved tree mortality rates ranging from 20% to 100%. Our results suggested a significant negative relationship between the amount of fire‐caused tree mortality and future dwarf mistletoe severity. These findings supported the presumed natural role of fire in altering dwarf mistletoe populations, which perhaps exhibits a linear relationship between fire‐caused host tree mortality and future dwarf mistletoe severity.     

Epidemiology of Heterobasidion abietinum and Viscum album on silver fir (Abies alba) stands of the Pyrenees

In the last two decades, stand decline and increased mortality has affected silver fir (Abies alba) forests in the Spanish Pyrenees. Simultaneously severe occurrences of the root rot fungus Heterobasidion annosum s.l. and of the mistletoe Viscum album have been reported. We aimed to improve the understanding of the epidemiology of both pathogens in our region. All H. annosum isolates found on silver fir were typed as H. abietinum. H. abietinum was more frequently observed where cuttings had targeted fir trees rather than other species. H. abietinum fruiting bodies were observed in the most recently cut stumps. V. album was more abundant on more dominant fir trees, and in southern aspect stands. The number of V. album colonies in the stand correlated (R² = 0.40) with silver fir mortality. Stands with a high level of V. album infection tended to have a smaller percentage of basal area in species other than silver fir, and they tended to be located on more south‐facing slopes. H. abietinum was widespread in silver fir forests of the Pyrenees. Our data suggest that, in the Pyrenees, the observed H. abietinum incidence may represent a combination of both primary and secondary spread of the pathogen. Favouring mixed forests should be tested as a potential control method for V. album. The correlation between silver fir mortality and V. album infection warrants further study, as the observed tree mortality could have occurred due to other factors than V. album, such as drought damage.     

Development of Pinus sylvestris Main Stems Following Three Different Precommercial Thinning Methods in a Mixed Stand

Reducing competition by point cleaning may be an attractive alternative to conventional precommercial thinning in forestry. In this study, survival and development of main stems of Scots pine (Pinus sylvestris L.) following point cleaning were examined in a mixed stand of mainly Scots pine and birch (Betula pendula Roth and Betula pubescens Ehrh.) in northern Sweden. Treatments studied were: no precommercial thinning, conventional precommercial thinning, point cleaning of all secondary stems, and point cleaning of only the secondary stems that were higher than the main stem, within a radius of 0.8 m from the main stem, respectively. A randomised complete block design with three replicates was used. Measurements taken 13 growing seasons after these treatments showed that conventional precommercial thinning was the best treatment for retaining Scots pines as main stems. However, conventional precommercial thinning resulted in the poorest timber quality of the remaining Scots pines in terms of stem taper, thickest branch diameter and relative live crown height. In conclusion, point cleaning has to be followed by a complementary cleaning operation if the objective is to retain Scots pines as main stems in such mixed stands.     

The effect of density after precommercial thinning on volume and structure in Pinus Sylvestris and Picea Abies stands

The interaction between stand density and dominant height and the development of volume, mean diameter, mortality and distribution of volume during a period from precommercial thinning to first commercial thinning was studied on permanent plots in Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) stands in Sweden. The reciprocal equation of the yield density effect was used to level the volume yield. Higher density after precommercial thinning resulted in higher yield and smaller mean diameter. The mortality up to first thinning was low, but is also dependent on density after precommercial thinning. The positive skewness of the volume distribution was higher in denser stands.     

Susceptibility of Coulter pine (Pinus coulteri) to western dwarf mistletoe (Arceuthobium campylopodum) in southern California

The susceptibility of Coulter pine (Pinus coulteri) to western dwarf mistletoe (Arceuthobium campylopodum) was evaluated in southern California. Eight stands infested with western dwarf mistletoe and dominated by Coulter and Jeffrey pines (Pinus jeffreyi) were sampled. Infection of Coulter pine was evaluated using circular plots established around large, severely infected Jeffrey pines and with rectangular transects. A total of 25 plots and 10 transects were sampled. In each plot and transect, live trees > 5 cm in diameter at breast height (1.4 m above the ground) were assigned a dwarf mistletoe rating (6‐class system). Nearly 600 Jeffrey pines and more than 400 Coulter pines were evaluated. Infection of Jeffrey and Coulter pines in the plots and transects averaged 96% and 97%, respectively. These data indicate that Coulter pine should be classified as a principal host of WDM and not as a secondary host as previously reported.     

Transformation of western hemlock (Tsuga heterophylla) tree crowns by dwarf mistletoe (Arceuthobium tsugense,Viscaceae)

Dwarf mistletoes (Arceuthobium species) are arboreal, hemiparasitic plants of conifers that can change the structure and function of the tree crown. Hemlock dwarf mistletoe (Arceuthobium tsugense subsp. tsugense) principally parasitizes western hemlock (Tsuga heterophylla) and effects 10.8% of all western hemlock trees in Oregon, USA. In this study, we climbed 16 western hemlock trees (age 97–321 years, height 33–54.7 m) across a gradient of infection (0%–100% of branches infected) and measured occurrence of all dwarf mistletoe infections, dwarf mistletoe caused deformities, foliage, branch and crown metrics, and sapwood area. We then modelled over 25 different response variables using linear and generalized linear models with three metrics of severity as explanatory variables: total infection incidence, proportion of all live branches infected, and proportion of all live, infected branches with 33 per cent or more foliage distal to infection. A strong effect of dwarf mistletoe intensification was the reduction of branch foliage and an increase in the proportional amount of foliage distal to infections, with severely infected trees having the majority of foliage distal to infections. Increasing severity led to an apparent crown compaction as crown volumes decreased and became increasingly comprised of deformities. Sapwood area was unrelated to infection severity. Branch length and diameters were unrelated to increasing infection severity despite severely infected branches supporting 1–70 infections. The most severely infected tree had 3,615 individual plants in the crown. Our results suggested that shifts in crown structure and branch deformation, foliage amount, and foliage distal to infection, reflected a likely reduction of capacity for tree growth that coincided with a hypothesized increase in resource demand by dwarf mistletoe plants as infection severity intensified.     

Forest thinning and subsequent bark beetle-caused mortality in Northeastern California

The Warner Mountains of northeastern California on the Modoc National Forest experienced a high incidence of tree mortality (2001–2007) that was associated with drought and bark beetle (Coleoptera: Curculionidae, Scolytinae) attack. Various silvicultural thinning treatments were implemented prior to this period of tree mortality to reduce stand density and increase residual tree growth and vigor. Our study: (1) compared bark beetle-caused conifer mortality in forested areas thinned from 1985 to 1998 to similar, non-thinned areas and (2) identified site, stand and individual tree characteristics associated with conifer mortality. We sampled ponderosa pine (Pinus ponderosa var ponderosa Dougl. ex Laws.) and Jeffrey pine (Pinus jeffreyi Grev. and Balf.) trees in pre-commercially thinned and non-thinned plantations and ponderosa pine and white fir (Abies concolor var lowiana Gordon) in mixed conifer forests that were commercially thinned, salvage-thinned, and non-thinned. Clusters of five plots (1/50th ha) and four transects (20.1 × 100.6 m) were sampled to estimate stand, site and tree mortality characteristics. A total of 20 pre-commercially thinned and 13 non-thinned plantation plot clusters as well as 20 commercially thinned, 20 salvage-thinned and 20 non-thinned mixed conifer plot clusters were established. Plantation and mixed conifer data were analyzed separately. In ponderosa pine plantations, mountain pine beetle (Dendroctonus ponderosae Hopkins) (MPB) caused greater density of mortality (trees ha⁻¹ killed) in non-thinned (median 16.1 trees ha⁻¹) compared to the pre-commercially thinned (1.2 trees ha⁻¹) stands. Percent mortality (trees ha⁻¹ killed/trees ha⁻¹ host available) was less in the pre-commercially thinned (median 0.5%) compared to the non-thinned (5.0%) plantation stands. In mixed conifer areas, fir engraver beetles (Scolytus ventralis LeConte) (FEN) caused greater density of white fir mortality in non-thinned (least square mean 44.5 trees ha⁻¹) compared to the commercially thinned (23.8 trees ha⁻¹) and salvage-thinned stands (16.4 trees ha⁻¹). Percent mortality did not differ between commercially thinned (least square mean 12.6%), salvage-thinned (11.0%), and non-thinned (13.1%) mixed conifer stands. Thus, FEN-caused mortality occurred in direct proportion to the density of available white fir. In plantations, density of MPB-caused mortality was associated with treatment and tree density of all species. In mixed conifer areas, density of FEN-caused mortality had a positive association with white fir density and a curvilinear association with elevation.     

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.     

抚育间伐对杉木人工林生长及出材量的影响

间伐对人工林生长及木材质量具有重要影响。在浙江开化开展了间伐强度（强度、中度和对照）和间伐方式（间伐1次,间伐2次）对杉木人工林生长及出材量等的影响研究,第1试验区大径材培育林分在第7年和第14年进行了2次间伐,第2试验区中径材培育林分仅在第12年时进行1次间伐。结果表明：22年生大径材培育林分和19年生中径材培育林分不同间伐处理之间树高、总断面积的差异均不显著。间伐显著增加了林木胸径的生长,大径材培育林分强度和中度间伐处理的平均胸径分别比对照增加13.78%和9.69%,中径材培育林分强度和中度间伐处理的平均胸径分别比对照增加12.28%和7.02%。间伐显著促进了林木单株材积的增加,2组试验林分活立木材积随着间伐强度的增加而降低,第1试验区22年生林分不同间伐强度之间活立木材积的差异不显著（P>0.05）,第2试验区强度间伐活立木材积显著低于对照（P<0.05）,中度间伐林分活立木材积与对照和强度间伐之间的差异不显著（P>0.05）。2组试验林分总蓄积和出材量也随着间伐强度的增加而降低,二者在3种间伐处理之间均没有显著差异（P>0.05）。间伐强度不能有效增加林分活立木材积和林分出材量,间伐次数对林分出材量及出材规格有重要影响。试验区杉木大径材培育以2次间伐,总间伐强度50%左右较为适宜,而中径材培育则以1次中度间伐（约25%）为宜。     

Effects of Mexican dwarf mistletoe (Arceuthobium vaginatum subsp. vaginatum) on the growth of Pinus cooperi in Durango,Mexico—A case study

Mexican dwarf mistletoe (Arceuthobium vaginatum subsp. vaginatum, Viscaceae) is the most widespread and damaging parasitic plant in Mexico. It parasitizes 10 species of pines (Pinus spp., Pinaceae) as principal hosts, including Pinus cooperi, one of the economically most important pines in the state of Durango. As a case study, we used stem analysis to estimate the effects of Mexican dwarf mistletoe on volume and height growth of dwarf mistletoe‐infected P. cooperi in western Durango (Ejido El Brillante, Municipality Pueblo Nuevo). This case study sampled a total of 48 trees, 12 in each of four infection classes estimated using the 6‐class dwarf mistletoe rating system (DMR): uninfected (DMR 0), lightly infected (DMR 1–2), moderately infected (DMR 3–4) and severely infected (DMR 5–6). Significant reductions in both volume and height growth were found for moderately and severely infected trees when compared to uninfected trees. On average, reductions in volume growth and height growth were as high as 50% and 17%, respectively. The largest growth reductions were for moderately infected trees, but large growth reductions also occurred for severely infected trees. Because of the reduced growth associated with moderate to severe infection, Mexican dwarf mistletoe‐infested pine forests in Durango, where timber production is a high priority, should be managed using harvesting practices that reduce dwarf mistletoe infection, and thereby, increase forest productivity.     

Canopy density in stands of Picea abies and Pinus sylvestris after different thinning methods

Altogether 82 plots (261 estimations) of Picea abies (L.) Karst, and 193 plots (360 estimations) of Pinus sylvestris (L.) stands were estimated by a vertical tube. The “crown free projection”, CFP, of stands thinned in three methods with different thinning grades was measured: unthinned, heavily and very heavily thinned, heavily thinned delayed first thinning, extra heavily thinned and thinned from the top. Basal area (m²ha^?1) density (stems ha^?1) and diameter sum (m ha^?1) were plotted against CFP. Basal area was the best practical measure of stand in this study. Generally Scots pine stands have higher CFP and the curves are steeper than in Norway spruce stands. Depending on the grade of thinning, heavily and very heavily thinned spruce stands, delayed first thinning included, have CFP values of 10–15% and stands thinned from the top, 20–40%, compared with 30–80% and 30–60% respectively in pine stands. Extra heavily thinned stands have the highest CFP, 20–80% in spruce and 50–90% in pine stands. The CFP levels after thinning are too high in pine stands for avoidance of sucker and sprout production of aspen and birch. In dense Norway spruce stands thinned from the top or heavily and very heavily thinned, the CFP values are low enough (≤30%) to diminish the production of suckers.     

Effects of thinning on plant species diversity and composition of understory herbs in a larch plantation

The effects of thinning on plant species diversity and composition of understory herbs in a larch plantation were investigated. The relationships between plant species diversity and composition of understory herbs and light conditions were established. Twenty-five 1 m × 1 m plots and fifteen 13 m × 1 m transects were set up in unthinned and thinned stands, respectively. All the transects in the thinned stands were set across the thinned rows and unthinned rows, and each of them was divided into nine 1 m × 1 m sub-plots. The herb diversity and light conditions were observed in each plot and sub-plot. The results show that there was a significant difference in herb diversity between the thinned and unthinned stands. All biodiversity indices except for evenness index in the thinned stand were higher than those of the unthinned stand, i.e., the herb diversity increased after thinning. According to the changes in herb densities and whether one species could be found in a stand or not before and after thinning, all herb species were classed into three types: positive, neutral and negative species, which referred to a species newly appeared and having an obviously increased density after thinning, with no obvious changes in its appearance and density after thinning, and disappeared and having an obviously decreased density after thinning, respectively. Many new species were found in the thinned stand like Corydalis pallida, Prenanthes tatarinowii, Vicia unijuga and Sonchus brachyotus etc. However, most species found in both the thinned and unthinned stands were negative species. In all nine subplots, only 11 and 10 species were found in spring and in autumn respectively, accounting for 17.74% and 15.15% of all the species in the thinned stand, respectively. All biodiversity indices were the highest in the center sub-plots and most of them tended to reduce from middle to side sub-plots. There was a close correlation between most of the three types of species and light conditions which was similar to each other in the thinned and unthinned stands. __________ Translated from Chinese Journal of Ecology, 2006, 25(10): 1201–1207 [译自: 生态学杂志]     

Silviculture as a disturbance regime: the effects of clear-cutting,planting and thinning on polypore communities in mixed forests

The structure of modern forest landscapes is profoundly affected by human-caused disturbances, particularly forest management; however, the effects and prospects of individual silvicultural techniques are insufficiently understood. This study distinguishes the effects of clear-cutting, planting and thinning on species richness and community composition of polypore fungi. In 2008–2009, 181 forest compartments (ranging from naturally regenerated deciduous stands to planted Picea abies stands and 0–137 years post clear-cutting) were explored in a hemiboreal landscape subjected to even-aged management. Altogether 104 polypore species were recorded. For species richness, time since clear-cutting was the most influential factor at both stand and landscape scales, followed by thinning. Clear-cuts had distinct polypore communities (including several red-listed species) whose species richness declined in time. Following 20 years post clear-cutting, species richness started to increase along different community–composition pathways determined by regeneration type. The communities developed after planting were moderately species rich at stand scale but homogeneous over larger areas. Thus, at landscape scale, mature unmanaged naturally regenerated stands hosted most species; thinning reduced species richness by approximately 15%, and among thinned stands, planted areas had a further 9–22% fewer species than naturally regenerated areas. In such variably managed landscape, silviculture appeared to create particularly distinct communities in young stands on nutrient-rich soils, which naturally provide polypores with a rich supply of small deciduous snags absent from stands artificially planted with P. abies and intensively thinned.     

The effects of heavy thinning on stem quality and timber properties of silver birch (Betula pendula Roth)

A study was carried out into the effects of heavy thinning onstem quality and timber properties of silver birch (Betula pendulaRoth). Comparisons were made with lightly thinned and unthinnedbirch stands. Birch stands from the UK were also compared withsimilarly managed stands from Germany and Denmark. In general,trees from lightly thinned and unthinned stands had similarcharacteristics. Heavy thinning enhanced diameter incrementthrough greater ring width in comparison with the other treatments.The timber properties of basic density, grain angle and shrinkagecoefficients in the radial, tangential and longitudinal directions,however, were largely unaffected. More and bigger branches andgreater stem taper were found on trees from heavily thinnedstands, but these stems still met the current Finnish veneerbirch quality and dimension standards. Comparisons between birchstands, managed by heavy and light thinning, from the UK andGermany and Denmark revealed no meaningful differences in timberproperties. The results suggest that veneer dimension and qualitylogs could be produced in the UK in rotations as short as 40years on good sites if heavy thinning is adopted.     

Commercial thinning that maintained species diversity of a mixed black spruce–jack pine stand enhanced productivity

Commercial thinning enables forest managers to meet timber production objectives. Thinning reduces tree density to alleviate competition for resources and favour growth of selected tree species. However, in doing so, thinning can homogenize the composition of mixed-species forests and raise biodiversity issues. There is increasing evidence that species richness can lead to higher productivity through a complementarity effect. Hence, thinning that would maintain species diversity of mixed-species forests could enhance stand productivity and help forest managers to reconcile timber production objectives and biodiversity issues. The objective of this study was to compare post-thinning stand production, experimentally over 10 years, in mixed and monospecific stands of black spruce (Picea mariana [Mill.] B.S.P.) and jack pine (Pinus banksiana Lamb.). The post-thinning stand production curve of the mixed stand converged toward that of the unthinned mixed stand while the production curves of the thinned and unthinned monospecific stands remained parallel. The convergent productivity of the mixed stand could be explained by a positive interaction between effects of thinning and niche complementarity. We propose that thinning that maintains species diversity of mixed stands could help forest managers who are implementing ecosystem management to reconcile timber production objectives with biodiversity issues.     

Effects of site preparation on emergence and early establishment of Larix olgensis in montane regions of northeastern China

Larix olgensis is one of the important commercial timber tree species in northeastern China, but it does not naturally regenerate in L. olgensis forests even after cutting. To identify the causes of the regeneration failure, the authors tested the effects of site preparation in a thinned stand and in a clearcut on the emergence and early establishment of L. olgensis seedlings. The results indicated that there was no correlation between cumulative emergence percentage (CEP) of L. olgensis seedlings and canopy openness, but the CEP was significantly affected by site preparation (P < 0.001). The average seedling survival rates after treatments consisting of removing both litter cover and understory vegetation cover were significantly higher in the clearcut than in the thinned stand (P < 0.001). Without site preparation, all seedlings in both the thinned stand and the clearcut disappeared within the first growing season, meaning that ground cover (understory vegetation cover and litter cover) was not beneficial to seedling survival. The average growth of seedlings exhibited significant differences between the thinned stand and the clearcut, and among the site preparations (P < 0.001). Intensely thinned stands without ground cover improved seedling growth. These results imply that light and ground cover might be the main factors limiting the early establishment of L. olgensis seedlings in managed plantation forests     

Impact of silviculture on dead wood and on the distribution and frequency of tree microhabitats in montane beech-fir forests of the Pyrenees

In forest ecosystems, the level of biodiversity is strongly linked to dead wood and tree microhabitats. To evaluate the influence of current forest management on the availability of dead wood and on the abundance and distribution of microhabitats, we studied the volume and diversity of dead wood objects and the distribution and frequency of cavities, dendrothelms, cracks, bark losses and sporophores of saproxylic fungi in montane beech-fir stands. We compared stands unmanaged for 50 or 100 years with continuously managed stands. A total of 1,204 live trees and 460 dead wood objects were observed. Total dead wood volume, snag volume and microhabitat diversity were lower in the managed stands, but the total number of microhabitats per ha was not significantly different between managed and unmanaged stands. Cavities were always the most frequent microhabitat and cracks the least frequent. Dendrothelm and bark loss were favored by management. Beech (Fagus sylvatica) carried many more microhabitats than silver fir (Abies alba), especially cavities, dendrothelms and bark losses. Fir very scarcely formed dendrothelms. Secondary tree species played an important role by providing cracks and bark losses. The proportion of microhabitat-bearing trees increased dramatically above circumference thresholds of 225 cm for beech and 215 cm for fir. Firs with a circumference of less than 135 cm did not carry microhabitats. In order to conserve microhabitat-providing trees and to increase the volume of dead wood in managed stands, we recommend conserving trees that finish their natural cycle over 10–20% of the surface area.