Drought sensitivity of Norway spruce is higher than that of silver fir along an altitudinal gradient in southwestern Germany

Context

For Central Europe, climate projections foresee an increase in temperature combined with decreasing summer precipitation, resulting in drier conditions during the growing season. This might negatively affect forest growth, especially at sites that are already water-limited, i.e., at low elevation. At higher altitudes trees might profit from increasing temperatures.

Aims

We analyzed variations in radial growth of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) along an altitudinal gradient from 400 until 1,140 m a.s.l. in the Black Forest, to assess climate responses with increasing elevation.

Methods

Climate–growth relationships were analyzed retrospectively using tree-ring and climate data. In total, we sampled stem discs of 135 trees to build 27 species- and site-specific chronologies (n _fir?=?13, n _spruce?=?14).

Results

Our results indicate distinct differences in climate–growth relations between fir and spruce along the gradient. Growth of high-altitude fir was positively related to temperature from January till March. Growth of low-altitude fir and spruce at all elevations was positively related to precipitation and negatively to temperature during the growing season, particularly in July. A self-calibrating Palmer drought severity index underlined summer drought sensitivity of these trees.

Conclusion

Overall, we found that climatic control of tree growth changes over altitude for fir. For spruce, a remarkable synchrony in growth variation and climate response was shown, which indicates that this species is drought sensitive at all studied elevations. In a future warmer climate, the growth of low-altitude fir and spruce along the entire studied gradient may be negatively affected in the Black Forest, if an increased evaporative demand cannot be compensated by increased water supply.     

Effect of seed source elevation on bud opening of Yezo spruce (<Emphasis Type="Italic">Picea jezoensis</Emphasis>)

Spring frost damage is one of the obstacles to be overcome in establishing a Yezo spruce (Picea jezoensis Carr.) plantation. The timing of bud opening of Yezo spruce seedlings cultured from seeds collected at three different elevations (420, 700, and 1200m) in the Tokyo University Forest in Hokkaido was observed at two planting site elevations (610 and 750m) to investigate whether seed source elevation or planting site elevation affects the bud opening date. At both planting sites the bud opening dates were not significantly different among the seedling groups from different seed source elevations. On the other hand, bud opening dates of seedlings planted at the lower elevation site were significantly earlier than those planted in the higher elevation site. It could be deduced from this study that the environment of the planting site rather than seed source elevation affects the bud opening date of Yezo spruce.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

峨眉山不同海拔冷杉径向生长对气候变化的响应

【目的】研究峨眉山不同海拔冷杉Abies fabri径向生长对气候变化的响应规律。【方法】以冷杉为研究对象,用树轮生态学的方法对峨眉山不同海拔的冷杉进行年轮采样、处理与分析,比较不同海拔冷杉径向生长特征及其与各气候因子之间的关系。【结果】年表统计数据表明,4个年表均具有较高的可靠性,可以用于树木生长与气候关系的研究。不同海拔冷杉径向生长与气候的关系具有明显的海拔差异,海拔2 400 m处冷杉径向生长与当年7月的温度呈显著正相关;海拔2 800 m处冷杉径向生长与当年4-9月的帕尔默干旱指数(PDSI)呈显著正相关;海拔2 900 m处冷杉径向生长与前一年9月和当年4月的温度呈显著负相关,与当年1-9月的PDSI均呈显著正相关;海拔3 000 m处冷杉径向生长和当年3-4月温度呈显著负相关,与当年6-7月的PDSI和当年4月平均相对湿度呈显著正相关。【结论】峨眉山高海拔的冷杉包含有更多的环境信息,对气候变化的响应也更为敏感;高海拔区域冷杉径向生长受到水分的强烈限制,而低海拔区域冷杉径向生长主要受温度影响;冷杉衰亡与气候因子的变化密切相关,随着区域气候的变暖,高海拔地区冷杉可能会进一步衰退。该研究揭示了峨眉山不同海拔冷杉生长动态及其对气候变化响应的敏感性差异,为全球气候变化背景下冷杉林的保护与适应性管理提供科学参考。     

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.     

Winter water relations of New England conifers and factors influencing their upper elevational limits. I. Measurements

The upper elevational limits of tree species are thought to be controlled by abiotic factors such as temperature and the soil and atmospheric conditions affecting plant water status. We measured relative water contents (RWC), water potentials (Psi) and cuticular conductances (g(c)) of shoots of four conifer species-eastern hemlock (Tsuga canadensis (L.) Carr.), eastern white pine (Pinus strobus L.), red pine (P. resinosa Ait.) and red spruce (Picea rubens Sarg.)-during two winters on Mt. Ascutney, Vermont, USA. Some micrometeorological measurements are also reported. Eastern hemlock and white pine were studied near their upper elevational limits at a 640-m site, and red pine was studied near its upper elevational limit at 715 m. Red spruce was also studied at the 715-m site, which is in the middle of its elevational range on this mountain. There was no evidence of winter desiccation stress in any species. The observed distribution of seedlings suggested that the upper elevational limits on shade-intolerant eastern white pine and red pine are set by the absence of suitable seed beds after 100 years without fire. Eastern hemlock is able to reproduce in deep shade on organic substrates, but germination at high elevations may be restricted by low temperatures.     

Species differences in total and vertical distribution of branch- and tree-level leaf area for the five primary conifer species in Maine, USA

Vertical distribution of leaf area largely governs both tree structure and function. Models of this important tree attribute have been constructed for several commercially important conifers. However, a limited number of studies have compared alternative modeling techniques and inherent species differences. This study used several existing datasets for the five primary conifer species in Maine, namely balsam fir [Abies balsamea (L.) Mill.], northern white-cedar [Thuja occidentalis (L.)], eastern hemlock [Tsuga canadensis (L.) Carr.], eastern white pine [Pinus strobus (L.)], and red spruce [Picea rubens (Sarg.)] to examine species variation in total and vertical distribution of projected leaf area at the individual branch- and tree-levels. In addition, multiple methods for modeling the vertical distribution of leaf area were examined across the species. For a given branch diameter and location within the crown, eastern hemlock branches held the greatest amount of leaf area, followed by balsam fir, northern white-cedar, white pine, and red spruce. At the tree-level, eastern white pine held the greatest amount of leaf area followed by eastern hemlock, balsam fir, red spruce, and northern white-cedar for a given tree size. Across species, the two-parameter, right-truncated Weibull distribution performed the best for predicting vertical distribution of leaf area when compared to the four-parameter beta and Johnson's S_B distributions (reduction of root mean square error of 1.7–21.1%). Northern white-cedar had a relative distribution of leaf area distinctly different than other species in this study with a mode shifted towards the upper crown. In contrast to red spruce and white pine, the mode of the relative distribution of leaf area for balsam fir and eastern hemlock occurred lower in the crown. Results of this study suggest that differences in total and vertical distribution of leaf area exist between species, but significant amounts of their variation are largely accounted for by bole and crown size.     

Seasonal changes in shoot water relations of Picea rubens at two high elevation sites in the Smoky Mountains

Seasonal changes in water relations of current-year shoots of red spruce (Picea rubens Sarg.) were examined in relation to climatic conditions in trees growing at elevations of 1720 and 1935 m on Clingman's Dome, Tennessee, USA, where increment core data have shown that red spruce decline increases with elevation. Relative height growth of trees at 1720 m was 68% greater than in trees at 1935 m. Following two weeks in July with only traces of precipitation, trees at both sites showed decreased saturated osmotic potentials. The magnitude of the reduction was greater in trees at the high elevation site than in trees at the low elevation site. However, during August and September, shoot water relations of trees at both sites were similar. Precipitation patterns and water relations measurements suggested that, at both sites, trees experienced water stress only briefly during the growing season and to a degree that could not account for the lower growth rates of trees at the high elevation site. During the period of cold hardening in October and November, trees at the low elevation site exhibited saturated osmotic potentials that were lower by 0.2 MPa and solute accumulation (osmol kg(dw) (-1)) that was 48% greater than in trees at the high elevation site.     

Effects of metals and pH on in vitro growth of Armillaria ostoyae and other root and butt rot fungi of red spruce

Armillaria ostoyae, Perenniporia subacida, Resinicium bicolor and Scytinostroma galactinum, root and butt rot fungi found on red spruce, Picea rubens, were tested, in vitro, for their sensitivity to metals typically found in high elevation forest soils where red spruce grows. Rhizomorph production by A. ostoyae from woody inocula in soils from red spruce stands at three elevations at each of five mountainous sites in the eastern United States was inhibited completely in the mineral soil from all elevations at all sites, and was also reduced significantly in the organic horizon from the upper two elevations at three of the sites. Inhibition was correlated with concentrations of metal ions in the soil. Growth of rhizomorphs into an agar medium containing lead and other heavy metals was inhibited for isolates of A. ostoyae from red spruce, but not for an isolate of Armillaria gallica from sugar maple; aluminium inhibited rhizomorph growth of isolates of both species. Mycelial growth of all four root and butt rot fungi was inhibited by lead, aluminium and other heavy metals depending on the solubility and concentration of metal and pH of the medium; growth inhibition was usually greater at an initial pH of 3.5 than at pH 4.5. Metal ions inhibited radial growth of Armillaria species more than that of the other three fungi. Rhizomorph growth of Armillaria was inhibited more than radial growth. Because local spread of A. ostoyae occurs frequently by means of rhizomorph growth between near roots, increases in lead, aluminium and other metals in the forest floor may contribute to this fungus' scarcity in high elevation soils and reduced incidence of infection at these sites in the eastern United States.     

Effects of shade on the morphology and physiology of amabilis fir and western hemlock seedlings

Successful regeneration of coastal montane sites harvested using alternative silvicultural systems may depend on the degree to which tree species can acclimate morphologically and physiologically to a variety of light environments. In a study to determine shade acclimation in montane conifers, one-year-old amabilis fir (Abies amabilis (Dougl.) Forbes) and western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings were grown in a nursery under four shade treatments: full sunlight (0% shade), 60% and 30% shade using shade cloth, and 30% shade using lath slats. Shading influenced shoot development, foliar physiology and morphological characteristics of both amabilis fir and western hemlock but in general, the effects were small. Shade levels of 60% were required to induce significant acclimation, and western hemlock appeared to respond more positively than amabilis fir and therefore was considered more shade tolerant than amabilis fir. Light quality had little influence on growth and development, as indicated by a lack of significant differences in physiology or morphology between seedlings grown under shade cloth or lath slats. There were indications that adequate nutrition levels may mitigate the effects of shade on seedling morphology and physiology.     

Evidence that longer needle retention of spruce and pine populations at high elevations and high latitudes is largely a phenotypic response

There is abundant evidence that evergreen conifers living at high elevations or at high latitudes have longer-lived needles than trees of the same species living elsewhere. This pattern is likely caused by the influence of low temperature in combination with related factors such as a short growing season and low nutrient availability. Because it is not known to what degree such patterns result from phenotypic versus genotypic variation, we evaluated needle longevity for common-garden-grown lowland populations of European Scots pine (Pinus sylvestris L.) of wide latitudinal origin and Norway spruce (Picea abies L.) of wide elevational origin. Nine-year-old trees of 16 Scots pine populations ranging in origin from 47 degrees to 60 degrees N were studied in Kórnik, Poland (52 degrees N) and 18-year-old trees of 18 Norway spruce populations ranging in origin from 670 to 1235 m elevation in southwestern Poland were studied near Morawina, Poland (51 degrees N, 180 m elevation). There was no tendency in either species for populations from northern or high elevation origins to retain needles longer than other populations. All of the Scots pine populations had between 2.5 to 3.0 needle age cohorts and all of the Norway spruce populations had between 6.4 and 7.2 needle age cohorts. Thus, extended needle retention in Scots pine and Norway spruce populations in low-temperature habitats at high elevations and high latitudes appears to be largely an environmentally regulated phenotypic acclimation.     

Long-term dynamics of a mixed conifer stand in Slovenia managed with a farmer selection system

The single-tree selection system is an important option for management of Norway spruce (Picea abies (L.) Karst.) and silver fir (Abies alba Mill.) forests because it provides continuous cover, requires low investments for tending, and promotes natural regeneration as well as high stand resistance and elasticity. It is often regarded as a very conservative system that usually results in only minor spatiotemporal changes in forest structure and composition. We studied management history, structural changes, regeneration dynamics, and light climate of a traditional single-tree farmer selection silver fir-Norway spruce forest (site typology Bazzanio-Abietetum). Stand structure was analyzed on five 0.25 ha permanent plots in 1994, 2001, and 2008. Regeneration density and height growth, forest floor vegetation, and light climate were also assessed on 1.5 × 1.5 m regeneration subplots in 2001 and 2008. Tree cores extracted from dominant trees from both species in two plots were used for reconstructing stand history and age structure of the canopy layer. We documented the forest response to three types of selection management regimes: excessive, normal, and conservative. Excessive management with harvest intensity significantly above the increment was documented until the late 1950s, including two peaks of heavy fellings (diameter limit cut) in the 1880s and 1930s, which favoured establishment of Norway spruce and released regeneration. The period that followed was characterized by normal selection management, but was nevertheless marked by a decline of silver fir as a result of air pollution and several droughts. This led to sanitary fellings that were carried out from the late 1970s to the early 1990s. In the last two decades conservative management followed, which led to suppression and decline of regeneration, especially of Norway spruce, and loss of selection structure. Although we recorded lower regeneration potential of silver fir compared with Norway spruce within the seedling category, silver fir outcompeted Norway spruce within the small-sized tree category (1 cm < dbh ? 10 cm) because of its superior height growth in low light levels (diffuse light <6%) and occupied a greater share of the canopy. Nevertheless, we anticipate that over the long-term the low light regime will also cause regeneration decline of silver fir and broadleaves. Our research revealed significant structural changes in a single-tree farmer selection forest during the last 150 years. These were a result of variable management regime and environment. A farmer single-tree selection system could better mimic the natural disturbance regime if spatiotemporal combinations of diverse felling regimes would be used.     

Effects of pocket gophers,bracken fern,and western coneflower on planted conifers in northern Idaho – an update and two more species

Grand Fir Mosaic forests innorthern Idaho are difficult to regenerate afternatural or human-caused disturbances. Sparseconifer regeneration appears to be associated withhigh populations of northern pocket gophers (Thomomys talpoides) plus expansion by bracken fern(Pteridium aquilinum (L.) Kuhn) and invasionof western coneflower (Rudbeckia occidentalisNutt.). This report updates an earlier study(Ferguson and Adams 1994) that quantified theeffects of four treatments on survival and growth ofplanted conifers: unweeded with gophers, weeded withgophers, unweeded without gophers, and weededwithout gophers. Weeding removed only bracken fernand western coneflower. Subalpine fir (Abieslasiocarpa (Hook.) Nutt.), grand fir (Abiesgrandis (Dougl.) Lindl.), and western larch (Larix occidentalis Nutt.) had over 80% mortalityfrom pocket gophers and other causes. Lodgepolepine (Pinus contorta var. latifoliaEngelm.) had 49.2% mortality from pocket gophersand 6.8% mortality from other causes, but snowdamage is a problem for lodgepole pine. Engelmannspruce (Picea engelmannii Parry ex Engelm.)had 42.2% gopher-caused mortality and 8.4%mortality from other causes. Spruce suffered littletop damage from snowpacks and was stout enough towithstand senescing bracken fern fronds. Westernwhite pine (Pinus monticola Dougl.) had only24.1% mortality from pocket gophers after 4 years,and only 11.8% of the white pine died from causesother than gophers. Of the mortality caused bypocket gophers, 76.8% occurred the first summer andnext two winters after planting. The recommendedspecies for reforestation are Engelmann spruce andwhite pine.     

Controls of growth phenology vary in seedlings of three, co-occurring ecologically distinct northern conifers

The objective of this study was to investigate the effects of temperature and seed-source elevation on height-growth phenology of three co-occurring and ecologically distinct northern conifers (Pinus contorta Dougl. ex Loud. var. latifolia (lodgepole pine), Picea glauca (Moench) Voss x Picea engelmannii Parry ex Engelm. (interior spruce) and Abies lasiocarpa (Hook.) Nutt. (subalpine fir)). Seed from populations of the three indigenous and co-occurring species was collected across an elevational transect on the southwestern slope of McBride Peak, near Prince George, BC. Collection sites were at elevations of 750 to 1850 m, the latter being close to the tree line. In 2003, seeds were germinated and seedlings raised under favorable growing conditions in a temperature-controlled glasshouse. In 2004, seedlings of each population were grown in natural daylengths at a location within 50 km of the seed collection site both in a temperature-controlled glasshouse and at a nearby field site, and height growth was recorded twice a week throughout the growing season. Species differed in both the date and the accumulated heat sum above 5 degrees C for the initiation and cessation of shoot extension. Growth durations (which integrate growth initiation and growth cessation) were more similar among species in the field than in the glasshouse. This suggests that different mechanisms of phenological control among co-occurring species can result in adaptive "equivalence" under a particular set of climatic conditions.     

A variance-covariance structure to take into account repeated measurements and heteroscedasticity in growth modeling

This study proposes a within-subject variance-covariance (VC) structure to take into account repeated measurements and heteroscedasticity in a context of growth modeling. The VC structure integrates a variance function and a continuous autoregressive covariance structure. It was tested on a nonlinear growth model parameterized with data from permanent sample plots. Using a stand-level approach, basal area growth was independently modeled for red spruce (Picea rubens Sarg.) and balsam fir [Abies balsamea (L.) Mill.] in mixed stands. For both species, the implementation of the VC structure significantly improved the maximum likelihood of the model. In both cases, it efficiently accounted for heteroscedasticity and autocorrelation, since the normalized residuals no longer exhibited departures from the assumptions of independent error terms with homogeneous variances. Moreover, compared with traditional nonlinear least squares (NLS) models, models parameterized with this VC structure may generate more accurate predictions when prior information is available. This case study demonstrates that the implementation of a VC structure may provide parameter estimates that are consistent with asymptotically unbiased variances in a context of nonlinear growth modeling using a stand-level approach. Since the variances are no longer biased, the hypothesis tests performed on the estimates are valid when the number of observations is large.     

Influence of slope position, stand type and rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech (Fagus orientalis Lipsky.) and spruce [Picea orientalis (L.) Link]

Oriental beech (Fagus orientalis Lipsky.) and Oriental spruce [Picea orientalis (L.) Link] are the two most common tree species in northeast Turkey. Their distribution, stand type and understorey species are known to be influenced by topographical landforms. However, little information is available as to how these changes affect litter decomposition rates of these two species. Here, we investigated the effects of slope positions (top 1,800 m, middle 1,500 m and bottom 1,200 m), stand type (pure and mixed stands) and purple-flowered rhododendron (Rhododendron ponticum) on litter decomposition rates of Oriental beech and spruce for 4 years using the litterbag technique in the field. Among these three factors, stand type had the strongest influence on litter decomposition (P < 0.001, F = 58.8), followed by rhododendron (P < 0.001, F = 46.8) and slope position (P < 0.05, F = 11.6). Litter decomposition was highest under mixed beech/spruce forest, followed by pure beech and spruce forest. Beech and spruce litter decomposed much faster in mixed bags (beech–spruce) than they did separately under each stand type. Purple-flowered rhododendron significantly reduced litter decomposition of Oriental beech and spruce. Beech and spruce litter decomposed much slower at top slope position than at either bottom or middle position. Differential litter decomposition of Oriental beech and spruce was mainly due to adverse conditions in spruce forest and the presence of rhododendron on the ground which was associated with lower soil pH. Higher elevations (top slope position) slowed down litter decomposition by changing environmental conditions, most probably by decreasing temperature as also other factors are different (pH, precipitation) and no detailed investigations were made to differentiate these factors. The adverse conditions for litter decomposition in spruce forest can be effectively counteracted by admixture of beech to spruce monoculture and by using the clear-cutting method for controlling rhododendron.     

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.     

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.     

Effects of overstory retention and site preparation on growth of planted white spruce seedlings in deciduous and coniferous dominated boreal plains mixedwoods

Survival and growth of planted white spruce was assessed under partial harvest treatments and different site preparation techniques in mixedwood forests of two compositions prior to logging: deciduous dominated (d-dom) – primarily comprised of mature trembling aspen (Populus tremuloides Michx.) and coniferous dominated (c-dom) – primarily comprised of mature white spruce (Picea glauca (Moench) Voss). Levels of overstory retention were 0% (clearcut), 50% and 75% of original basal area, and site preparation techniques were inverted mounding, high speed mixing, scalping and control (no treatment). The survival and growth of white spruce were assessed seven years after planting. The experiment was established as a part of the Ecosystem Management Emulating Natural Disturbance (EMEND) experiment located in northern Alberta, Canada. In the c-dom, the 50% and 75% retention of overstory resulted in reduced growth and survival of white spruce seedlings compared to clearcuts. In contrast, in the d-dom, the seedlings performed best in sites that had 50% of the overstory retained. For the c-dom, the mounding and mixing treatments yielded the best growth of spruce seedlings, while scalping yielded the worst. In the d-dom, spruce growth was highest in sites with the mixing treatment. In the d-dom, growth and survival of the planted spruce was greater than in the c-dom. The natural regeneration of deciduous trees was suppressed by the retention of canopy regardless of original composition.     

Canopy disturbance and intertree competition: implications for tree growth and recruitment in two yellow birch–conifer stands in Quebec, Canada

Composition, structure, and species-specific patterns of recruitment and growth were characterized in two yellow birch (Betula alleghaniensis Britt.)–conifer stands in Quebec, Canada, to improve our understanding of the dynamics of these complex ecosystems. The mixture of mid- and shade-tolerant species in the canopy, the inverse J-shape stem diameter distribution, and the age distribution were indicative that the two stands were in a late-successional stage. Recruitment of mid-tolerant species above 1.3 m in height appeared to be periodic and synchronized with historical spruce budworm (Choristoneura fumiferana Clem.) outbreaks, while the coniferous component of these mixedwood stands recruited continuously. Results suggest that recruitment of yellow birch and red maple (Acer rubrum L.) requires disturbances of a certain intensity that affect at least 25 % of the forest cover. In contrast, balsam fir (Abies balsamea (L.) Mill.) and red spruce (Picea rubens Sarg.) can recruit under the canopy without relying on moderate or large canopy disturbances. Results suggest that the historical disturbance regime, and differences in shade tolerance between species, largely govern the contemporary composition of these stands. This study improves the comprehension of mechanisms that regulate the dynamics of yellow birch-conifer stands and will be useful for the subsequent elaboration of forest management strategies.