Estimating cumulative defoliation of balsam fir from hemlock looper and balsam fir sawfly using aerial defoliation survey in western Newfoundland,Canada

Both hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) and balsam fir sawfly (Neodiprion abietis (Harris)) undergo periodic outbreaks in eastern Canada and cause significant growth and mortality losses to forests. Tree growth and mortality are closely related to cumulative defoliation estimates, which integrate annual defoliation over multiple years. Our objective was to determine a method to estimate cumulative defoliation of balsam fir (Abies balsamea [L.] Mill) due to these insects in western Newfoundland, using aerial defoliation survey data, as an essential input to modeling impacts for Decision Support Systems. Interpretation of aerial defoliation survey data for hemlock looper and balsam fir sawfly is problematic because both insects feed upon multiple age classes of foliage. Current-year (2008) aerial defoliation survey data were compared with ground estimates of defoliation by age class from 45 plots (450 trees and 395 mid-crown branch samples), representing a range of defoliation severity classes for each insect. Cumulative defoliation was calculated using defoliation per foliage age class, weighted by relative foliage mass for a given age of foliage. Three significantly different severity classes were defined based on cumulative defoliation values derived from aerial defoliation survey: (i) 1-year moderate (30–70%) defoliation, (ii) 1-year severe (71–100%) defoliation with calculated cumulative defoliation values of 19 and 39%, respectively, for balsam fir sawfly, 21 and 34% respectively for hemlock looper; and (iii) 2–3 years of moderate–severe defoliation, with cumulative defoliation ranging between 59 and 64% for balsam fir sawfly and 49% for hemlock looper. Defoliation severity from aerial defoliation survey alone hence can be misleading if defoliation measurements are not converted to cumulative defoliation values.     

Broom rusts of balsam fir and black spruce in Newfoundland

The broom rusts of balsam fir and black spruce occur sporadically throughout the island of Newfoundland, but they are not a serious threat to the forests. The incidence and intensity of the rusts vary, but no tree mortality can be attributed to the diseases. The average number of brooms per tree was higher in black spruce than in balsam fir. Also, more brooms occurred on brandies than on tree trunks. Height and diameter growth was less in infected trees than in uninfected trees of both the species.     

Snag dynamics in post-harvest landscapes of western Newfoundland balsam fir-dominated boreal forests

We examined changes in standing dead tree (snag) density and biomass with time following harvest across a chronosequence of balsam fir (Abies balsamea) dominated boreal forests in western Newfoundland, Canada. Current snag management practices in Newfoundland recommend a minimum of 10 snags/ha on the post-harvest landscape. Snags declined significantly in the first two decades of the chronosequence. The rapid rate of decline in snag density which occurred immediately post harvest was likely attributable to windfall and domestic harvest for firewood. A second rapid rate of snag density decline occurred 10–15 years post harvest which potentially reflected the average lifespan of snags in western Newfoundland. Average snag densities approximated the minimum management goal during the period from 15 to 60 years since harvest (YSH). However, 53–60% of sites sampled in the 15–60 YSH period contained <10 snags/ha. Snag densities then increased with forest age, again reaching high levels 81–100 YSH which were comparable to the density at the beginning of the chronosequence. Achieving the goal of 10 snags/ha on all post-harvest sites in western Newfoundland, especially between 15 and 60 YSH, will require changes to current forest management practices.     

An effect of gravity on bud-burst in balsam fir

Bud-burst on first order lateral branches of Abies bafsamea L. (balsam fir) was delayed when the branches were rotated 180 degrees about their long axis. This was not a consequence of injury caused by the treatment because buds rotated 180 degrees on inverted plants flushed at the same time as the controls, whereas flushing of all other buds was delayed. Buds thus appear to be more vigorous when maintained in the same orientation to gravity in which they are formed and the site of gravitational stimulus perception appears to be the bud itself. Except on the leading shoot, leaves from inverted buds turned so that their adaxial surface faced upward, unless there was intense illumination from below. However, both anisophylly and positioning of leaves on lateral shoots were apparently predetermined because the shorter, more forward pointing leaves appeared below the longer distichous leaves on shoots from inverted buds. Shoots with normally oriented leaves appeared the next season.     

Impacts of hemlock looper defoliation on growth and survival of balsam fir, black spruce and white birch in Newfoundland, Canada

Hemlock looper (Lambdina fiscellaria fiscellaria (Guen.)) is an economically damaging defoliator that undergoes periodic outbreaks in Newfoundland, Canada. It defoliates and causes extensive tree mortality to its primary host, balsam fir (Abies balsamea [L.] Mill.). We quantified tree survival using data from permanent sample plots (PSPs) and growth reduction or release using dendrochronology, and related these impacts to defoliation severity determined from annual aerial defoliation survey data. Such impact relationships are necessary as a fundamental input to a Decision Support System. Growth and survival of balsam fir, black spruce (Picea mariana (Mill.) B.S.P.) and white birch (Betula papyrifera Marsh.) were assessed from 1996 to 2008 in 48 Newfoundland Forest Service PSPs, selected based on four classes of defoliation severity. Two years of severe (71-100%) defoliation resulted in almost complete mortality for balsam fir, 10 years after defoliation, whereas survival was 70-80% for black spruce and white birch. Lower defoliation severity (1-2 years of moderate (31-70%) or 1 year of severe) resulted in approximately 60% survival for balsam fir and no reduction in survival for black spruce and white birch. Maximum growth reduction of balsam fir was 10-15% with 1 year of moderate-severe defoliation, 35-40% with 2 years of moderate defoliation, and about 50% with 2 years of severe defoliation. Growth recovered to pre-defoliation rates 5 years after defoliation ceased in all severity classes. Growth reduction and recovery of black spruce were more variable and lower than for balsam fir, and white birch exhibited only minor (<10%) growth reduction during the defoliation year or 1 year after defoliation. Control measures should focus on avoiding severe defoliation for two consecutive years.     

Response of balsam fir and hemlock roots to injuries

Drill wounds in balsam fir and hemlock roots activated the nonspecific resistance mechanisms of compartmentalization in wood and necrophylactic periderm in bark. Tangential bands of resin ducts localized around the wounds constituted the barrier zones in the secondary xylem of conifer roots. Barrier zones were more extensive in roots which showed symptoms characteristic of invasion by fungi and bacteria after wounding. This observation supports an expanded definition of barrier zones; barrier zones may form not only in response to mechanical wounds but also in response to xylem injury caused by pathogens. Multiple bands of resin ducts were common in young xylem when bark lesions developed around wounds. Necrophylactic periderms isolared dead bark from living bark. Development of phellem cells with dark contents and thick suberized walls, typical of exophylactic periderm, followed initiation of necrophylactic periderm. The wound responses were similar in both balsam fir and hemlock.     

Certain seasonal changes in balsam fir cambium and its derivatives

Summary The seasonal cambial activity of balsam fir producing normal and compression wood was observed. Samples were taken throughout one growing season. Activity was initiated in normal wood with the precocious development of one sieve cell per file 2 weeks before activity was initiated in compression wood with the formation of one new sieve cell per file. Deactivation dates in normal and compression wood were the same.Dating of the phloem adjacent to normal wood can be carried out using crushed sieve cells or an annually formed band of tannin filled parenchyma whereas adjacent to compression wood, only crushed sieve cells can be used.The maximum cambial zone widths in normal and compression wood were 15 and 16 cells, respectively. The final xylem to phloem ratio in normal wood was 14:1 while in compression wood it was 21:1.Time required for development of tracheids was examined throughout the growing season. Growth trends for the radial growth and differentiation phases were studied in both normal and compression wood. Tracheid development in normal wood took between 14 and 33 days while in compression wood it varied between 5 and 49 days.Research financed under McIntire-Stennis Research Project 5009 and COE Projects R625-173 and R625-195     

Separation of eastern spruce and balsam fir by chemical methods

Summary Increased utilization of eastern spruce and balsam fir has led to a need for a quick method of separating these woods in a mill situation. One such method might be the use of a chemical indicator. A test of various classes of chemical agents applied at different seasons of the year showed that a pH indicator might be suitable for achieving a separation. The most suitable indicator was tested on samples from different geographic locations and at three highproduction stud mills. Additional tests were conducted to explore such variables as moisture content and surface condition of the wood, type of solvent, concentration and temperature. The most suitable indicator found was bromophenol blue at a concentration of 0.10 percent in 95 percent ethanol. When applied to green wood which had been allowed to dry for a few minutes to a few hours, this indicator produced various shades of orange, yellow, green or blue with spruce and a dark blue or blue-violet with fir. With an understanding of the variables that affect the reaction, it is felt that bromophenol blue can be used for the separation of eastern spruce and balsam fir on a commercial basis.The authors would like to thank fellow researchers in New York, Vermont, New Hampshire and New Brunswick, Canada, for providing samples for testing. Also, assistance from the following stud mills in Maine is greatly appreciated: Diamond International Corporation (Passadumkeag), Georgia-Pacific Corporation (Woodland) and St. Regis Paper Company (Costigan). This research was financed under McIntire-Stennis Research Project 5-5-39602.     

A comparison of red spruce and balsam fir shoot structures

I compared the shoot structures of high-elevation red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea (L.) Mill.). Needle widths, thicknesses and perimeters were measured to estimate total leaf areas from measured projected leaf areas. Measured needle perimeter/needle width ratios differed significantly from estimated ratios that assumed needles were either rhomboidal or elliptical in cross section. The vertical and horizontal silhouette shoot area to total leaf area ratios (STAR(v) and STAR(h)) of the two species were negatively correlated with needle packing and canopy height. Red spruce had higher values of STAR(v) than balsam fir at each canopy height, but STAR(v) declined with canopy height at a similar rate in the two species. The STAR(h) values of the two species did not differ significantly at a given canopy height. Needle packing increased with canopy height at the same rate in the two species. Needle weight increased in red spruce and decreased in balsam fir with increased needle packing, but showed no significant dependence on canopy height. Red spruce had higher values of STAR(h) than balsam fir at low values of needle packing, but STAR(h) values converged at high values of needle packing. The generally comparable values of STAR, along with similar needle diameters, may imply that red spruce and balsam fir have similar collection efficiencies of wet and dry particles. Measurements of STAR may be used to estimate leaf area indices (LAI) more accurately when using indirect techniques.     

Carbon and biomass partitioning in balsam fir (Abies balsamea)

Balsam fir (Abies balsamea (L.) Mill) was extensively sampled to investigate the effects of forest management practices, site location, within-crown position, tree component (i.e., stem, foliage, branches and roots), and tree social classes on biomass and carbon (C) partitioning at the individual tree level and across ecological regions. The sites were located in three ecologically distinct forest regions of west-central New Brunswick, Canada. There were no significant differences in %C content of trees across ecological regions or across tree social classes. However, at the individual tree level, significant differences were evident in biomass and C allocation between different parts of the tree, between treatment types (i.e., unmanaged and pre-commercially thinned stands) and between within-crown positions, indicating the need for separate estimates of biomass and C content of tree components to obtain more precise estimates of quantities at the stand level. Calculating stand C content based on constant allocation values, as is commonly done, produced errors of up to 15% compared with the values calculated in this study. Three allometric equations of biomass and C that account for partitioning among different parts of the tree were developed and compared: (1) a third-order polynomial, (2) a modified inverse polynomial and (3) a modified Weibull equation. Diameter at breast height (DBH) was used as the only explanatory variable to describe fresh biomass, dry biomass and C content. All regressions derived showed a high correlation with DBH, with most r2 values > 0.95. A comparison of the equation results showed that the modified Weibull equation gave consistent results with the best overall fit and was the simplest of the three equations investigated. The regressions can be used to estimate forest biomass and tree C content at the stand level, given specific information on DBH.     

Springtime resumption of photosynthesis in balsam fir (Abies balsamea)

Photosynthesis in balsam fir (Abies balsamea (L.) Mill.) was measured in the field at two locations in New Brunswick, Canada from late winter to late spring in 2004 and 2005. No photosynthesis was detectable while the soil remained below 0 degrees C throughout the rooting zone. In both years, photosynthesis began once soil temperature rose to 0 degrees C. In potted seedlings in growth chambers, there was no photosynthesis at an air temperature of 10 degrees C if the pots were frozen. These findings suggest that, once air temperatures permit photosynthesis, it is the availability of unfrozen soil water that triggers the onset of photosynthesis. In the field, full recovery of photosynthetic capacity following the onset of soil thaw was dependent on air temperature and took 5 weeks in 2005, but 10 weeks in 2004. There were two substantial frost events during the recovery period in 2004 that may explain the extended recovery period. In 2005, recovery was complete after the accumulation of 200 growing degree days above 0 degrees C after the start of soil thaw.     

Changes in the water relations of balsam fir and white birch saplings after thinning

In the autumn of 1987, young balsam fir (Abies balsamea (L.) Mill.) and white birch (Betula papyrifera Marsh.) trees were thinned and their water relations followed during the next two growing seasons. At the beginning of the first summer following treatment, thinned trees of both species had lower osmotic potentials at full saturation (Psi(pi,sat)) and at turgor loss point (Psi(pi,tlp)) compared with controls. At this time, Psi(pi,sat) was linearly related to the percentage of full sunlight reaching the trees. A higher sugar concentration in leaves was an important component of the lower Psi(pi,sat) of thinned trees. For the other two sampling dates during the first growing season after treatment and all three sampling dates during the second growing season after treatment, little osmotic adjustment of the thinned trees relative to the control tress was observed in either species. The absence of osmotic adjustment during the second growing season following thinning suggests that other mechanisms were responsible for the acclimation of the treated trees to the higher atmospheric evaporative demand. Sapwood permeability (k) of white birch was higher than that of balsam fir, but no differences in k or in sapwood area were found between treated and control trees of either species. Predawn water potentials (Psi(pred)) of treated trees were less negative than those of controls.     

A method for reconstructing the development of the sapwood area of balsam fir

Leaf area is commonly estimated as a function of sapwood area. However, because sapwood changes to heartwood over time, it has not previously been possible to reconstruct either the sapwood area or the leaf area of older trees into the past. In this study, we report a method for reconstructing the development of the sapwood area of dominant and codominant balsam fir (Abies balsamea (L.) Mill.). The technique is based on establishing a species-specific relationship between the number of annual growth rings in the sapwood area and tree age. Because the number of annual growth rings in the sapwood of balsam fir at a given age was found to be independent of site quality and stand density, the number of rings in sapwood (NRS) can be predicted from the age of a tree thus: NRS = 14.818 (1 - e(-0.031 age)), unweighted R(2) = 0.80, and NRS = 2.490 (1 - e(-0.038 age)), unweighted R(2) = 0.64, for measurements at breast height and at the base of the live crown, respectively. These nonlinear asymptotic regression models based only on age, were not improved by adding other tree variables such as diameter at breast height, diameter at the base of the live crown, total tree height or percent live crown.     

Relationships of climate and cell features in stems and roots of black spruce and balsam fir

? The anatomical differences of mature black spruces and balsam firs were examined at stem and root level in order to characterize their wood properties at cellular level and link these differences to climate.

? Anatomical variability of these species was evaluated in relation to climate data gathered from 2001 to 2004 during the cell enlargement (CE) and wall thickening and lignification (WTL) phases. Lumen area, single cell wall thickness and total tracheid radial diameter were analyzed and regrouped into earlywood and latewood.

? Results from a principal component analysis (PCA) indicated that both first eigenvectors account for 82% and 90% of total variance for CE and WTL respectively. These component factors revealed that precipitation, humidity and number of days with precipitation significantly influence the lumen area (p = 0.0168) and radial cell diameter (p = 0.0222) in earlywood. Significant differences were registered between species and tree parts (stem and root) for the lumen area, radial cell diameter and cell wall thickness in both earlywood and latewood.

? In our study, black spruce exhibited smaller tracheid size in both stem and roots compared to balsam fir. Furthermore, the lower amount of tracheids produced during the growing season and higher proportion of latewood ensure a higher wood density of black spruce. The influence of temperature on earlywood formation is significant, whereas no influence was observed on latewood.

     

Seasonal changes in soil and tissue nutrition in balsam fir and influence on postharvest needle abscission

Postharvest needle abscission in balsam fir varies based on time of harvest, but little is known about the effect of soil and tree nutrition on postharvest needle abscission. From April to November, 18 balsam fir samples were obtained each month and evaluated for postharvest needle abscission characteristics. Soil samples were collected simultaneously and evaluated for moisture, pH, organic matter, cation exchange capacity, and mineral nutrition. Needle tissue samples were also evaluated for mineral nutrition. Many parameters were found to vary based on month throughout the study, but not all were associated with needle abscission. Soil organic matter, cation exchange capacity, Ca, and Mn all had a significant (P?P?R²?>?.50), suggesting that higher concentrations of tissue Ca were linked to superior needle retention in balsam fir. Overall, this study provides evidence that changes to soil management, such as the inclusion of organic matter and certain minerals, may improve needle retention in balsam fir.     

Differences in stem respiration responses to temperature between balsam fir trees in thinned and unthinned stands

The change in stem respiration rates with change of temperature was significantly lower in a recently thinned balsam fir (Abies balsamea (L.) Mill.) stand than in an unthinned stand. A comparison of respiration measurements made in the morning with those made in the afternoon indicated that there was more variation among afternoon readings and greater responses to changes in temperature among morning readings. Lower respiration rates in the afternoon were more common in the thinned stand than in the unthinned stand.     

Post-wildfire seeding in forests of the western United States: An evidence-based review

Broadcast seeding is one of the most widely used post-wildfire emergency response treatments intended to reduce soil erosion, increase vegetative ground cover, and minimize establishment and spread of non-native plant species. We conducted an evidence-based review to examine the effectiveness and effects of post-wildfire seeding treatments on soil stabilization, non-native species invasion, and plant community recovery in the western U.S. We reviewed 94 scientific papers and agency monitoring reports identified using a systematic search protocol. As sampling designs have become more rigorous in recent years, evidence that seeding is effective in reducing erosion has decreased. Of highest and high quality studies evaluating soil erosion, 92% (11 of 12) were published since 2000, none of which showed an effective result. Before 2000, the majority of the studies (71%) fell into the lowest quality categories, of which 72% showed seeding to be effective. The majority of studies (20 of 27, 74%) evaluating soil erosion in seeded versus unseeded controls showed that seeding did not reduce erosion relative to unseeded controls. Even when seeding significantly increased vegetative cover, seeded sites rarely supported plant cover levels considered sufficient to stabilize soils within the first and second year post-wildfire. Of the 11 studies evaluating seeding effectiveness for curtailing invasions of non-native plant species, an almost equal percentage found seeding treatments to be either effective (54%, 6 studies) or ineffective (45%, 5 studies). However, the majority of effective and ineffective treatments (83% and 80%, respectively) used non-native species. Sixteen of 26 studies (62%) evaluating seeding effects on plant communities reported that seeding suppressed recovery of native plants, although data on long-term impacts of this reduction are limited. The literature suggests that post-wildfire seeding does little to protect soil in the short-term, has equivocal effect on invasion of non-native species, and can have negative effects on native vegetation recovery, although long-term studies are needed to assess lasting impacts of seeded species.     

Inter-tree and intra-tree variations in ring width and wood density components in balsam fir (Abies balsamea)

This study quantified and compared intra-tree and inter-tree variations in ring width and wood density components in balsam fir (Abies balsamea) grown in Quebec, Canada. In addition, the study examined correlations between ring width and wood density components at different stem positions from the stump level to the stem top. Ring width and wood density components of individual rings were measured by X-ray densitometry. Both the intra- and inter-tree variations in balsam fir are large, but the inter-tree variation is relatively smaller than the intra-tree variation. Much of the intra-tree variation is due to the radial variation, whereas the axial variation is much smaller. Compared to ring width and its components, wood density characteristics show a considerably smaller variation at both the inter- and intra-tree level. In almost all wood characteristics studied (except for latewood width), the intra- and inter-tree variations are more or less influenced by tree age. Cambial age explains more intra-tree variation in wood density components than ring width, whereas more intra-tree variation in ring width components is due to ring width. Cambial age and ring width explain a comparable percentage of variation in ring density. Only a few of the correlations between ring width and wood density components vary significantly with stem position from the stump to the stem top at the inter-tree level. In balsam fir, a negative correlation between ring density and ring width is significant in the butt log, but the correlation decreases to an insignificant level at and above a height of 3.0 m.     

Ecophysiology and growth of advance red spruce and balsam fir regeneration after partial cutting in yellow birch-conifer stands

We investigated ecophysiological and growth responses of short (0.4 to 1.3 m in height) advance regeneration of red spruce (Picea rubens Sarg.) and balsam fir (Abies balsamea L.) six years after removal of 0, 40, 50, 60 and 100% of the overstory basal area (BA) in two yellow birch-conifer stands. Partial cuts significantly increased stomatal conductance of red spruce only. Light-saturated photosynthesis (leaf-area basis) of both species increased with BA removal, but unlike red spruce, specific leaf area (SLA) of balsam fir decreased with increased cutting intensity. Partial cuts appreciably increased the concentration of N and Ca in red spruce and balsam fir foliage, respectively, and resulted in decreased foliar concentrations of K in red spruce and Mg in balsam fir. The height and lateral growth of both species increased with BA removal, although partial cuts were more beneficial to balsam fir. The data suggest that short advance regeneration of red spruce and balsam fir can coexist under partial overstory conditions, but balsam fir has physiological characteristics and a capacity for morphological adjustment (SLA) that places it at an advantage when in competition with red spruce.