Susceptibility of ponderosa pine, Pinus ponderosa (Dougl. ex Laws.), to mountain pine beetle, Dendroctonus ponderosae Hopkins,attack in uneven-aged stands in the Black Hills of South Dakota and Wyoming USA

Mountain pine beetle, Dendroctonus ponderosae Hopkins can cause extensive tree mortality in ponderosa pine, Pinus ponderosa Dougl. ex Laws., forests in the Black Hills of South Dakota and Wyoming. Most studies that have examined stand susceptibility to mountain pine beetle have been conducted in even-aged stands. Land managers increasingly practice uneven-aged management. We established 84 clusters of four plots, one where bark beetle-caused mortality was present and three uninfested plots. For all plot trees we recorded species, tree diameter, and crown position and for ponderosa pine whether they were killed or infested by mountain pine beetle. Elevation, slope, and aspect were also recorded. We used classification trees to model the likelihood of bark beetle attack based on plot and site variables. The probability of individual tree attack within the infested plots was estimated using logistic regression. Basal area of ponderosa pine in trees ≥25.4 cm in diameter at breast height (dbh) and ponderosa pine stand density index were correlated with mountain pine beetle attack. Regression trees and linear regression indicated that the amount of observed tree mortality was associated with initial ponderosa pine basal area and ponderosa pine stand density index. Infested stands had higher total and ponderosa pine basal area, total and ponderosa pine stand density index, and ponderosa pine basal area in trees ≥25.4 cm dbh. The probability of individual tree attack within infested plots was positively correlated with tree diameter with ponderosa pine stand density index modifying the relationship. A tree of a given size was more likely to be attacked in a denser stand. We conclude that stands with higher ponderosa pine basal area in trees >25.4 cm and ponderosa pine stand density index are correlated with an increased likelihood of mountain pine beetle bark beetle attack. Information form this study will help forest managers in the identification of uneven-aged stands with a higher likelihood of bark beetle attack and expected levels of tree mortality.     

Effect of soil temperature on root and shoot growth and on budburst timing in conifer seedling transplants

Root growth in seedling transplants of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), Pacific silver fir (Abies amabilis(Dougl.) Forbes), noble fir (Abies procera Rehd.), lodgepole pine (Pinus contorta Dougl. ex Loud.) and ponderosa pine (Pines ponderosa Doug. ex Laws.) began when soil temperature exceeded 5 °C. Root growth increased rapidly after 10 °C and attained maximum values at 20 °C. At 30 °C, no root growth occurred in the firs; in the pines, root growth was 30 to 39% of maximum. Maximum shoot growth also occurred at 20 °C. In ponderosa pine, height growth of seedlings from a high-elevation source was unaffected by cold soil, but in low-elevation seedlings it was reduced. Budburst in Douglas-fir and the pines was delayed up to 11 days by cold soil, whereas in silver fir and noble fir, it was only slightly delayed. Prior to new root growth in ponderosa pine, xylem pressure potentials and stomatal conductances during the afternoon indicated reduced stomatal opening at all soil temperatures, whereas 23 days later, stomata were open to a greater degree when temperatures exceeded 10 °C.     

Yield reduction in spruce infected with Armillaria solidipes in the southern interior of British Columbia

The effect of non‐lethal root infection on the yield of planted spruce (Picea engelmannii x glauca hybrid spruce) in southern British Columbia was determined in a 34‐year‐old, mixed‐species stand infected with Armillaria solidipes Peck. Yield reduction correlated with trees’ infection duration. Individual trees infected for 24 years had about 42% less volume than their healthy equivalents. Using the incidence of infection in stands in this area and the individual losses per tree, reduction at the stand level was approximated at 11% and increasing.     

Modeling the probability of observing Armillaria root disease in the Black Hills

Armillaria root disease (ARD) occurrence on ponderosa pine (Pinus ponderosa) is not uniform in the Black Hills of South Dakota, USA. To help manage this ecosystem, a model was developed to predict the probability of observing diseased trees. A kernel density estimator was used to estimate the probability of observing ARD using presence data from two field studies. An eight‐parameter regression equation using topographical data (Universal Trans Mercator coordinates, elevation and slope) derived from a Digital Elevation Model was fitted to the estimated probabilities and the residuals kriged to produce correction factors for the regression estimates. The final model, which had a relative mean squared error of 0.128, identified two peaks of high probability in the north‐west portion of the Black Hills and several peaks of moderate probability throughout the Black Hills.     

Auxin-impregnated hygroscopic gel: effects on ponderosa pine and common hackberry seedlings

Greenhouse and field studies were conducted to determine the effects of indole3-butyric acid (IBA) and 2,4-dichlorophenoxyacetic acid (2,4-D) on root development and survival of 3+0 bareroot Pinus ponderosa (Dougl. ex Laws.) and 2+0 bareroot Celtis occidentalis (L) seedlings. In the greenhouse, 100 and 200 ppm. IBA and 10 and 50 ppm 2,4-D were applied to seedlings through a root dip in an auxin hygroscopic gel mix. A randomized complete block analysis of variance indicated that IBA gel treatments increased root volume and root dry weight in hackberry and had no effect on ponderosa pine; 2,4-D gel treatments had detrimental effects on both species. In the field, IBA gel treated ponderosa pine seedlings exhibited improved height and survival relative to the non-treated control seedlings. There was no detectable improvement in survival in the field for hackberry. However, diameter and stem dry weight were lower for hackberry seedlings treated with the gel dip alone.     

Growth of young Pinus ponderosa and Pinus contorta on compacted soil in central Washington

Growth in height, diameter, and volume was measured on 9- to 18-year-old ponderosa pine (Pinus ponderosa) and 10- to 13-year-old lodgepole pine (Pinus contorta) trees growing on or near compacted skid trails in the Yakima Indian Reservation in south-central Washington. Soil bulk density of the 0- to 30.5-cm deep layer was measured with a single-probe nuclear densimeter on two sides of each sample tree and in adjacent undisturbed soil. On three ponderos pine sites logged 23 years before the study, average bulk density on skid trails was 15% greater than on adjacent undisturbed soil. On a lodgepole pine site logged 14 years before the study, soil on skid trails averaged 28% greater bulk density than undisturbed soil.Total growth of ponderosa pine and the last 5 years of growth were significantly related (P = 0.07) by regression analysis to age of trees, site index, basal area of the adjacent overstory, and the percentage of increase in soil bulk density. At the mean increase in soil bulk density, total height, diameter, and volume growth were reduced 5%, 8%, and 20%, respectively.Total growth of lodgepole pine and the last 5 years of height, diameter, and volume growth were significantly related to tree age and the percentage of soil organic matter. Increase in soil bulk density was not significantly related to growth of this species.     

Correlation of seedling size and branch number with disease resistance of Pinus thunbergii seedlings to Bursaphelenchus xylophilus

Japanese black pine (Pinus thunbergii) seedlings resistant to pine wood nematode (PWN; Bursaphelenchus xylophilus) are routinely selected in Japanese field inoculation trials. Correlations between morphological factors (such as height, stem diameter at ground level and number of branches on seedlings) and disease resistance were examined to improve the production efficiency of 1‐year‐old black pine seedlings for inoculation. Family relatedness and environmental conditions strongly affected seedling resistance; accordingly, logistic regression analysis was used to separate effects of these two variables. Height and stem diameter at ground level significantly correlated with disease resistance in seedlings inoculated with PWN. Because (a) interactions between stem diameter at ground level and environmental condition were significant and (b) height did not interact with any other factor, it was concluded that height of 1‐year‐old Japanese black pine seedlings independently correlated with PWN resistance. Thus, field inoculation tests should use tall seedlings to achieve enhanced survival rates.     

Damage by pine weevil Hylobius abietis to conifer seedlings after shelterwood removal

Abstract

Pine weevil (Hylobius abietis L.) damage to seedlings after overstorey removal was investigated in a survey study in six shelterwoods in the south–central part of Sweden. The shelterwoods predominantly consisted of Scots pine, except at one site where the shelter trees mainly consisted of Norway spruce. Before final cutting, 10 plots were laid out at each site and measurements of shelter trees and marked seedlings were taken. The seedlings were examined during the 2 years after final cutting. The study showed that removal of shelter trees increases the risk of severe damage by pine weevil and the variable that was most strongly correlated with the risk was the seedling root collar diameter. Both Scots pine and Norway spruce seedlings were severely damaged by pine weevil, and most of the feeding occurred during the first year after cutting. The amount of debarked area was significantly larger for Scots pine than for Norway spruce seedlings. Vitality (growth of the leading shoot before final cutting) of the seedlings also affected the probability of damage. Seedlings with high vitality were less damaged by pine weevil than seedlings with low vitality. For Scots pine the shelterwood density before final cutting was correlated to the intensity of pine weevil feeding after cutting. In conclusion, after the final cutting of a pine or spruce shelterwood, pine weevils will probably invade the area. To avoid serious damage, Norway spruce and Scots pine seedlings should have reached a diameter of at least 10–12 mm.     

Container seedlings outperform barefoot stock: Survival and growth after 10 years

Ponderosa pine, Jeffrey pine, and Douglas-fir seedlings were planted in container or bareroot form at three elevations in northern California. At the lowest elevation (762 m), container seedlings of ponderosa pine were significantly taller than bareroot seedlings at ages 4 and 10, and had breast-height diameters that were significantly larger than bareroot counterparts at age 10. Survival of Douglas-fir container seedlings was significantly greater than that of barefoot seedlings for all ages tested. At the mid-elevation site (1220 m), container seedlings of ponderosa pine and Douglas-fir were significantly larger in breast-height diameter than bareroot seedlings at age 10. Douglas-fir container seedlings survived significantly better at all ages than barefoot seedlings. At the highest elevation (1662 m), seedling height and diameter did not differ significantly, but survival of container seedlings was significantly higher than barefoot seedlings for both pine species at all ages tested.     

Seasonal changes in above- and belowground carbohydrate concentrations of ponderosa pine along a pollution gradient

Seasonal patterns of carbohydrate concentration in coarse and fine roots, stem or bole, and foliage of ponderosa pine (Pinus ponderosa Laws) were described across five tree-age classes from seedlings to mature trees at an atmospherically clean site. Relative to all other tree-age classes, seedlings exhibited greater tissue carbohydrate concentration in stems and foliage, and greater shifts in the time at which maximum and minimum carbohydrate concentration occurred. To determine the effect of environmental stressors on tissue carbohydrate concentration, two tree-age classes (40-year-old and mature) were compared at three sites along a well-established, long-term O3 and N deposition gradient in the San Bernardino Mountains, California. Maximum carbohydrate concentration of 1-year-old needles declined with increasing pollution exposure in both tree-age classes. Maximum fine root monosaccharide concentration was depressed for both 40-year-old and mature trees at the most polluted site. Maximum coarse and fine root starch concentrations were significantly depressed at the most polluted site in mature trees. Maximum bole carbohydrate concentration of 40-year-old trees was greater for the two most polluted sites relative to the cleanest site: the bole appeared to be a storage organ at sites where high O3 and high N deposition decreased root biomass.     

Comparison of the virulence of Armillaria cepistipes and Armillaria ostoyae on four Norway spruce provenances

The basidiomycetes Armillaria cepistipes and Armillaria ostoyae frequently occur in the same forest stand. In this study, we determined the virulence of 20 isolates of A. cepistipes and 16 isolates of A. ostoyae on four different provenances of 2‐year‐old Norway spruce (Picea abies). Within 30 months after inoculation, 1.1 and 19.1% of the seedlings inoculated with A. cepistipes and A. ostoyae, respectively, had died or were dying. The incidence of dead and dying seedlings varied between 3 and 49% among the A. ostoyae isolates. The virulence of an isolate was positively correlated to its ability to produce rhizomorphs. One Norway spruce provenance showed significantly lower susceptibility to A. ostoyae than the other three. Rhizomorphs of both Armillaria species were attached to the root surface. The attached rhizomorphs of A. ostoyae, however, were associated with significantly more lesions. The virulence of the isolates was not correlated with their wood‐degrading capability for either of the Armillaria species.     

Morphological characteristics associated with tolerance to competition from herbaceous vegetation for seedlings of jack pine,black spruce,and white pine

Tolerance of bareroot and container-grown seedlings of black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), and eastern white pine (Pinus strobus L.) to competition from herbaceous vegetation was examined in the first five years after planting on a site in the Great Lakes/St. Lawrence forest of Ontario, Canada. Shoot and root morphological characteristics of various stocktypes were measured before planting and correlated with 5-year survival and growth following control and no control of herbaceous vegetation. For black spruce and jack pine, medium-sized bareroot stocktypes had greater relative 5-year stem volume growth in the presence of herbaceous vegetation than did container stock of either species or large bareroot stock of spruce. Relative volume growth was measured as the ratio of the cumulative stem volume increment in the presence of vegetation (Veg) to that in the absence of vegetation (NoVeg), i.e., the Veg:NoVeg ratio. In white pine, the Veg:NoVeg ratio of volume increment of medium container and large bareroot stocktypes exceeded that of small container and medium bareroot stocktypes. In jack pine, root collar diameter at planting and number of first-order lateral roots were positively correlated with 5-year Veg:NoVeg ratio of volume increment. In white pine, the Veg:NoVeg ratio was also positively correlated with root collar diameter at planting and with root volume. In black spruce, the ratio was not related to pre-plant morphology. Thus, for white pine and jack pine, certain pre-plant morphological features may be useful in forecasting the relative ability of different stocktypes to grow under herbaceous competition conditions in the field.     

Impact of armillaria root disease and the effect of thinning in a late‐rotation Pinus radiata plantation

In a search for alternative methods for managing armillaria root disease in Pinus radiata plantations in New Zealand, a thinning trial was established in a 1985 planted stand infested by Armillaria novae‐zelandiae. The incidence of chronic (non‐lethal) infection rose significantly following a first thinning at age 7 years that provided additional inoculum in the form of colonized stumps. However, this effect diminished before age 19 years when the small stumps had decomposed and, contrary to expectations, a further increase in infection did not follow a second thinning at age 13.5 years, despite a high incidence of larger colonized second thinning stumps. Apparently the increased vigour and associated resistance in the released older trees was sufficient to counter any increase in inoculum potential following treatment. It therefore now appears that thinning may not be detrimental to the final crop volume in contemporary diseased second‐rotation plantations. However, this treatment furnishes no disease control benefit, and should still be deployed judiciously in heavily infested stands. There was a small, but significant reduction in volume growth on the more severely infected trees during the first 13 years in this trial, but on the larger trees later in the rotation at age 19 years there was no difference in stem volume between infection severity categories. A previous assessment based only on the earlier data therefore overestimates the impact of Armillaria in present and future pine plantations in New Zealand. Nevertheless, its widespread distribution and its demonstrated pathogenicity indicate that forest owners should be conscious of the presence of Armillaria during plantation management.     

Hydraulic conductivity in roots of ponderosa pine infected with black-stain (Leptographium wageneri) or annosus (Heterobasidion annosum) root disease

Roots from healthy and diseased mature ponderosa pine, Pinus ponderosa Laws., trees were excavated from a site near Burns, Oregon. The diseased trees were infected with black-stain root disease, Leptographium wageneri Kendrick, or annosus root disease, Heterobasidion annosum (Fr.) Bref., or both. Axial hydraulic conductivity of the roots was measured under a positive head pressure of 5 kPa, and the conducting area was stained with safranin dye to determine specific conductivity (k(s)). In diseased roots, only 8-12% of the cross-sectional xylem area conducted water. Resin-soaked xylem completely restricted water transport and accounted for 13-16% of the loss in conducting area. In roots with black-stain root disease, 17% of the loss in conducting area was associated with unstained xylem, possibly resulting from occlusions or embolisms. Based on the entire cross-sectional area of infected roots, the k(s) of roots infected with black-stain root disease was 4.6% of that for healthy roots, whereas the k(s) of roots infected with annosus root disease was 2.6% of that for healthy roots. Although these low values were partly the result of the presence of a large number of diseased roots (72%) with no conducting xylem, the k(s) of functional xylem of diseased roots was only 33% of that for healthy roots. The low k(s) values of functional xylem in diseased roots may be caused by fungus induced occlusions preceding cavitation and embolism of tracheids. The k(s) of disease-free roots from diseased trees was only 70% of that for healthy roots from healthy trees. The disease-free roots had the same mean tracheid diameter and tissue density as the healthy roots, suggesting that the lower k(s) in disease-free roots of diseased trees may also have been caused by partial xylary occlusions.     

Results of a thinning study in a black stain (Leptographium wageneri)‐infested Jeffrey/ponderosa pine forest in northern California

Black stain root disease (BSRD) caused by Leptographium wageneri var. ponderosum (Verticicladiella wageneri) is a destructive disease in many Jeffrey pine (Pinus jeffreyi) and ponderosa pine (P. ponderosa) forests of the western United States. Overcrowding is a pervasive problem in these forests, unless alleviated by thinning. In an overcrowded BSRD‐infested second‐growth Jeffrey/ponderosa pine forest in northern California, a large study (16.2 hr; 40 acres) was designed to determine the impact of thinning to four levels—6.9 m²/ha (30 ft²/acre), 13.8 m²/ha (60 ft²/acre), and 20.7 m²/ha (90 ft²/acre) of basal area and a control with no thinning—on the incidence of BSRD‐caused mortality. Ten years after this experiment was initiated, results indicate that thinning to any of these levels lowered the incidence of BSRD‐caused mortality as compared with BSRD‐caused mortality in unthinned control plots.     

Physiological responses of ponderosa pine in western Montana to thinning, prescribed fire and burning season

Low-elevation ponderosa pine (Pinus ponderosa Dougl. ex. Laws.) forests of the northern Rocky Mountains historically experienced frequent low-intensity fires that maintained open uneven-aged stands. A century of fire exclusion has contributed to denser ponderosa pine forests with greater competition for resources, higher tree stress and greater risk of insect attack and stand-destroying fire. Active management intended to restore a semblance of the more sustainable historic stand structure and composition includes selective thinning and prescribed fire. However, little is known about the relative effects of these management practices on the physiological performance of ponderosa pine. We measured soil water and nitrogen availability, physiological performance and wood radial increment of second growth ponderosa pine trees at the Lick Creek Experimental Site in the Bitterroot National Forest, Montana, 8 and 9 years after the application of four treatments: thinning only; thinning followed by prescribed fire in the spring; thinning followed by prescribed fire in the fall; and untreated controls. Volumetric soil water content and resin capsule ammonium did not differ among treatments. Resin capsule nitrate in the control treatment was similar to that in all other treatments, although burned treatments had lower nitrate relative to the thinned-only treatment. Trees of similar size and canopy condition in the three thinned treatments (with and without fire) displayed higher leaf-area-based photosynthetic rate, stomatal conductance and mid-morning leaf water potential in June and July, and higher wood radial increment relative to trees in control units. Specific leaf area, mass-based leaf nitrogen content and carbon isotope discrimination did not vary among treatments. Our results suggest that, despite minimal differences in soil resource availability, trees in managed units where basal area was reduced had improved gas exchange and growth compared with trees in unmanaged units. Prescribed fire (either in the spring or in the fall) in addition to thinning, had no measurable effect on the mid-term physiological performance and wood growth of second growth ponderosa pine.     

Higher resistance of the offspring of Scots pine trees resulting from natural regeneration in old foci of Heterobasidion annosum root rot

Abstract

Heterobasidion annosum (Fr.) Bref. is a fungal pathogen causing annosum root rot – one of the most economically important diseases in coniferous stands. The major aim of this study was to compare the resistance of the offspring of Scots pine trees (Pinus sylvestris L.) from seed orchards and the offspring of trees that were naturally reproduced in old foci of the disease. In experiments conducted in vitro, we used 960 seedlings which were grown from seeds collected from 60 trees in 6 old foci of the disease and 640 seedlings grown from seeds collected in 4 seed orchards. The offspring of trees from seed orchards after inoculation with H. annosum had nearly twofold higher mortality rate than the seedlings developed from the seeds collected in old foci of the disease. This suggests that the offspring of self-sown trees in old foci of the disease has a greater, genetically conditioned resistance to annosum root rot.     

Tree vigour and the susceptibility of Douglas fir to Armillaria root disease

In this study the effects of thinning, fertilization and pruning on the vigour of Douglas fir (Pseudotsuga menziesii) and its susceptibility to Armillaria root disease were explored. Tree vigour was defined as the relative capacity for tree growth, expressed as the above-ground biomass increment per unit of photosynthetic tissue, or growth efficiency (GE). It has been hypothesized that trees with higher GE can better resist pathogen attack, and that GE can be used as a predictor of tree susceptibility to disease. In a previous study, four Douglas fir plantations were thinned, fertilized and pruned in all combinations, and the effects of these treatments on tree vigour were measured after 10 years. Root disease was not a factor in the initial study design, and mortality was ignored until 8 years after the treatments were applied. The results of an earlier study were utilized and the correlation between Armillaria root disease incidence and the effects of earlier stand treatments on tree growth was tested. Armillaria ostoyae was the primary mortality agent in the study area. Disease incidence on infested subplots ranged from 2 to 20%. Armillaria ostoyae incidence was the highest at medium tree density (6.1%), slightly lower on the low density (5.6%), and lowest on the unthinned plots (3.8%). There were no significant correlations between disease incidence and previous tree growth. The vigour of trees that became symptomatic or died by 1993 was not significantly different in 1983–85 from the vigour of trees that remained asymptomatic. On these sites, in areas of infestation, A. ostoyae is killing the largest, fastest growing trees, as well as less vigorous trees. Armillaria continues to cause mortality, regardless of the growth efficiency or growth rate of the host.     

Detection of Diplodia pinea in asymptomatic pine shoots and its relation to the Normalized Insolation index

Diplodia pinea (syn. Sphaeropsis sapinea), a common pathogenic fungus, causes considerable damage in Italy, particularly to pine stands in which trees are subjected to environmental stress. The occurrence of D. pinea in symptomless Pinus nigra shoots was investigated and related to the amount of radiation received by the trees growing on a site in a year, expressed as the Normalized Insolation index (NIi). Twenty‐seven pines were selected from nine locations in Trentino (northern Italy). For each pine the incidence of the fungus in apparently healthy shoots was determined by both culturing on an agar medium and application of real‐time PCR. The incidence of D. pinea determined by culturing samples taken from asymptomatic trees was 59% (16 of 27 trees), compared with 85% found using real‐time PCR (23 of 27 trees). Detection of the pathogen in healthy pine tissue was positively correlated (p < 0.05) with the NIi values, using both detection methods.     

Trichoderma atroviride promotes growth and enhances systemic resistance to Diplodia pinea in radiata pine (Pinus radiata) seedlings

Root drench application of Trichoderma atroviride isolates R32, R33, R40 and R84 promoted the growth of potted radiata pine seedlings. After 6 weeks, seedlings treated with R33 and R84 had thicker stems and greater stem and root biomass (p < 0.05) than untreated controls. Treatment with R32 increased seedling root biomass whilst R40 increased stem diameter. None of the isolates affected seedling height. One isolate, R33, induced systemic resistance to stem inoculation with Diplodia pinea and reduced dieback incidence by 20% compared with untreated controls. To our knowledge, this is the first report of systemic induced resistance by Trichoderma in a pine species. Furthermore, seedlings that were treated with R33 (root drench) plus foliar application of methyl jasmonate (MeJA) expressed elevated peroxidase activity in their stems 2 weeks later, compared with seedlings treated only with MeJA. Because R33 itself did not affect peroxidase activity, this may be indicative of treatment synergy or defence potentiation by R33. Curiously, R33 + MeJA induced terpenoids but suppressed phenylalanine ammonia‐lyase activity suggesting possible trade‐offs between phenolic and terpenoid defence pathways in the treated seedlings.