Size-mediated climate-growth relationships in temperate forests: A multi-species analysis

In most dendrochronological studies, climate-growth relationships are established on dominant trees to minimize non-climatic signals. However, response to environmental factors may be affected by tree-size, which begs the question of the representativeness of dominant trees on the stand level. To highlight the variations in climate-growth relationships among sizes and species, under a wide range of ecological conditions (climate and soil properties), 61 pure even-aged stands were sampled across France. At each stand, two tree-ring chronologies were established from 10 big- to 10 small-diameter trees. Our objectives were, (1) to assess variations in climate sensitivity between the two size-diameter classes, and (2) to investigate the role of species and ecological conditions on these variations. The climate-growth relationships were evaluated from 122 tree-ring chronologies (1 220 trees) through extreme growth years and correlation function analyses. Sensitivity to climate of shade-intolerant and moderately shade-tolerant species (Picea abies (L.) Karst., Pinus sylvestris L. and Quercus petraea (Matt.) Liebl.) remained constant between the size-diameter classes for both temperature and hydric balance, while the shade-tolerant species Abies alba Mill. and Fagus sylvatica L. displayed significant differences, with larger trees being more sensitive to summer drought than smaller trees. This difference increased with increasing climatic xericity. Our results suggest that, for shade-tolerant species, (1) big trees could be more sensitive to climatic change especially under xeric climate, and (2) future tree ring studies should include trees stratified by size to produce unbiased estimation of sensitivity to climate.     

农田防护林更新树种选择及配置模式研究

对农田防护林更新树种和配置模式进行了分析研究。从新品种的引种上筛选出适于河西走廊发展的农田防护林树种为三倍体毛白杨B10 0 8和BT85 ,并能在瘠薄和盐碱化土壤上良好生长 ;经抗寒性分析得出垂柳、云杉、白榆、柽柳、沙枣抗冻性较强 ;为减缓天牛等蛀干害虫危害 ,在林网配置模式上引入了云杉、侧柏、樟子松、垂柳、小叶白蜡、国槐等耐瘠薄抗寒性强的树种 ,营造多树种林带 ,组成混交林网结构 ,使林网中感虫杨树的比例占 2 0 %左右 ,增加了林网结构的稳定性和长效性     

Effects of enhanced nitrogen deposition on foliar chemistry and physiological processes of forest trees at the Bear Brook Watershed in Maine

Effects of enhanced nitrogen deposition on nutrient foliar concentrations and net photosynthesis of sugar maple (Acer saccharum Marsh.), American beech (Fagus grandifolia Ehrh) and red spruce (Picea rubens Sarg.) were evaluated at the Bear Brook Watershed in Maine (BBWM). The BBWM is a paired-watershed forest ecosystem study with one watershed treated since 1989 with bimonthly dry ammonium sulfate ((NH₄)₂SO₄) additions at a rate of 25.2 kg N ha⁻¹ year⁻¹, while the other watershed serves as a reference. The (NH₄)₂SO₄ treatment resulted in significant increases in foliar N concentrations for all three species and significant reductions in foliar Ca, Mg and Zn concentrations for American beech and red spruce. Treatment effects on foliar concentrations of other nutrients were not significant in any species. Despite higher N concentrations in all species, only treated sugar maple showed significantly higher photosynthetic rates. The non-response in net photosynthesis to higher foliar N in American beech and red spruce might be attributed to their low foliar Ca and/or Mg concentrations. Higher net photosynthetic rates in sugar maple might be explained by the higher foliar N and by the ability of this species to maintain an adequate Ca and Mg supply. Results suggested that nutrient imbalances due to inadequate supply of Ca and Mg might have counteracted a potential increase in net photosynthesis induced by higher N concentrations in American beech and red spruce.     

Forest regeneration 50 years following partial cutting in mixedwood ecosystems of southern Quebec,Canada

The impact of winter harvesting on regeneration 50 years after an experimental diameter-limit cutting was examined in mixed deciduous–coniferous ecosystems of southern Quebec, Canada. The study was conducted in La Mauricie National Park, Quebec, Canada. Regeneration data in two balsam fir (Abies balsamea (L.) Mill.), red spruce (Picea rubens Sarg.), sugar maple (Acer saccharum Marsh.), and yellow birch (Betula alleghaniensis Britt.) ecosystem types were analyzed. Comparisons between uncut and cut stands were obtained from a total of 63 sample plots. For both ecosystems, there were no significant differences between uncut and cut plots for regeneration density and stocking. The most abundant regeneration species were balsam fir, red spruce, sugar maple, red maple (Acer rubrum L.), yellow birch and American beech (Fagus grandifolia Ehrh.). The type of diameter-limit cutting described in the study did not affect regeneration density and stocking but its impact on productivity, timber quality and genetics is still unknown.     

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.     

Investigations of the pore-size distribution of wood in the dry and wet state by means of mercury intrusion porosimetry

This paper deals with the determination of the pore-size distribution of untreated and thermally modified twin samples of Norway spruce (Picea abies (L.) Karst.) and sycamore maple (Acer pseudoplatanus L.) by means of mercury intrusion porosimetry. The investigations considered oven-dried and at 22 °C and 95 % relative humidity-conditioned samples. A special sample holder and a novel method for evacuation were developed for the measurements. This was necessary both to ensure that samples were positioned at a defined distance and the conditioned samples did not dry in an uncontrolled manner. The results clearly show that the climatic state influences the results. Thermal modification greatly alters the pore-size distribution of spruce. This can be largely attributed to the formation of intercellular spaces in the middle lamella as a result of cell-wall compression due to thermal modification. The alteration of the pore structure of maple due to thermal modification is not as pronounced as that of spruce.     

Climatic signal in annual growth variation of silver fir (Abies alba Mill.) and spruce (Picea abies Karst.) from the French Permanent Plot Network (RENECOFOR)

This paper explores the growth/climate relationships in earlywood, latewood and total ring-width chronologies of five Norway spruce (Picea abies) and six silver fir (Abies alba) stands sampled in the French permanent plot network (RENECOFOR) (327 trees). The relationships between climate and ring widths were analyzed using extreme growth years, simple correlations and response functions analysis (bootstrapped coefficients). Monthly climatic regressors were derived by a physiological water balance model that used daily climatic data and stand parameters to estimate soil water deficits. Pointer years underline the high sensitivity to winter frosts (1956, 1986) and exceptional annual droughts (1962, 1976, 1991) for both species. For those years, growth variations were higher for Abies alba than for Picea abies. For each species, the climate information of tree ring series is not modified by local site characteristics (altitude, slope, aspect, soil water reserve). Moreover, strong specific differences appear among species. Earlywood and latewood spruce growth mainly depends on current summer soil water deficit conditions. For silver fir, winter and early summer temperatures, as well as the water supply of the previous year (August to October) play a major role for the production of earlywood, after which the current early summer water supply influences mainly latewood width.     

Models to predict time since death of Picea abies Snags

Number of years since death was estimated by dendrochronological cross-dating of 107 standing dead trees (snags) of Norway spruce [Picea abies (L.) Karst.] in a submountainous old-growth forest in south–central Norway. Snag characteristics (size, bark cover, branch order present and variables derived from tree-ring analyses) were used in stepwise linear regression procedures to identify variables that explained time since death. Number of branch orders present (where branches growing directly on the stem were branch order 1, branches growing on order 1 branches were order 2, and so on) explained two-thirds of the variation in time since death. Adding other significant variables, such as diameter, relative height of snags, percentage bark cover and average tree-ring width in the final years before death, increased model precision only moderately. The models were validated by the PRESS statistic, which showed that new observations were predicted fairly well with 65–69% of the variation explained.     

Temperature sensitivity and recruitment dynamics of Siberian larch (Larix sibirica) and Siberian spruce (Picea obovata) in northern Mongolia’s boreal forest

This study sought to clarify the recruitment dynamics and growth of Siberian larch (Larix sibirica) and Siberian spruce (Picea obovata) in relation to changing temperatures in northern Mongolia. These tree species are the primary forest species found in the closed-canopy boreal forest of north-central Mongolia. Mongolia’s boreal forests exist along the southern terminus of the Siberian boreal system in both pure and mixed species stands. I collected tree cores and cross-sections as well as site and tree stature parameters from 118 forest plots in the Darhad valley of north-central Mongolia. Principle components analysis of 130 L. sibirica tree ring series informed the construction of two composite chronologies for this species. A chronology for P. obovata was developed using 24 tree ring series. Correlation analysis between tree ring indices and temperature data showed two distinct growth signals: a positive response to growing season temperatures was exhibited by one L. sibirica chronology and a negative response to spring temperatures was exhibited by a second L. sibirica chronology. The P. obovata chronology exhibited strong negative correlations with mean monthly and mean maximum monthly growing season temperatures. Multiple analyses of variance (MANOVA) indicated that tree stature (dbh, height) and site parameters (latitude, longitude, slope, aspect, elevation) did not significantly predict growth response or species. Forest recruitment events appear episodic for both species. Synchronous establishment of saplings, based on approximate root collar age, suggests an initial floristic model for mixed composition stands likely due to supra-annual variations of fire, land-use and climate. Forest management activities in the region should consider the diverging growth response to temperature shown here by prioritizing protection forests and the various ecosystem services provided by forests in arid ecosystems. In addition, promoting selection harvests over clear-felling would maximize future alternatives under conditions of rapidly changing climate. Care should be taken in new forest management planning activities until adequate information exists on the likely trajectory of this system due to climate-induced forest change.     

Toward improved understanding of the cell-wall density and porosity of wood determined by gas pycnometry

The main objective of this study was to evaluate the effect of sample preparation on cell-wall density and porosity using gas pycnometry. Native and thermally modified twin samples of Norway spruce (Picea abies (L.) Karst.), sycamore maple (Acer pseudoplatanus L.), and European ash (Fraxinus excelsior L.) were analyzed. The samples differed in terms of shape, geometry, and climatic state. On the one hand, the samples were oven dry as usual and, on the other hand, conditioned at 22 °C and 95 % relative humidity. Furthermore, the samples were processed using solvent exchange drying. In addition to helium, nitrogen was used as a displacement gas. The tests show i.a. how this can lead to misinterpretation of the cell-wall density or porosity of wood determined by gas pycnometry. The results show that native spruce has a lower cell-wall density and higher porosity compared with native maple and ash. Due to thermal modification, the cell-wall densities are decreased. The investigations show that the determined cell-wall density and porosity of the wood are strongly dependent on the sample geometry and climatic state. The cell-wall densities of all investigated wood species in the conditioned state at 22 °C and 95 % relative humidity are significantly higher compared with the oven-dry cell-wall densities.     

New observation of cone flies attacking cones ofPicea obovata andLarix sibirica in central Siberia

Two species of cone flies (Strobilomyia anthracina andS. svenssoni) emerged from puparia were collected under Siberian spruce (Picea obovata) trees in the central part of the Krasnoyarsk region, central Siberia. Parasites of the Figitidae and Braconidae families emerged from 28.1% of the puparia, and fungal and bacterial diseases destroyed 24.8% of the puparia. Weight and length of the puparia depended on the sanitary status. Two types of cone fly eggs were observed on spruce cones. At least three species ofStrobilomyia (S. Infrequens, S. laricicola, and S. sibirica) attacked cones of Siberian larch (Larix sibirica) in central Siberia. (Responsible editor: Zhu Hong)     

The pollen record of a 20th century spruce beetle (Dendroctonus rufipennis) outbreak in a Colorado subalpine forest,USA

The frequency and intensity of ecosystem disturbance, including outbreaks of forest insects and forest fires, is expected to increase in the future as a result of higher temperatures and prolonged drought. While many studies have concentrated on the future climatic impacts on fire, little is known about the impact of future climate on insect infestation. Paleoecological techniques are important in this regard in identifying the potential relationships between climate and insect outbreaks in the past, as a predictive tool for the future. We examine a high-resolution 20th century record of spruce beetle (Dendroctonus rufipennis) infestation from a small, subalpine lake, comparing the paleoecological record to the historical and tree-ring record of the event. An extensive spruce beetle outbreak occurred in northwestern Colorado during the 1940s and 1950s, causing widespread mortality of mature Picea engelmannii. Pollen analysis of this period documents the decline of Picea and its replacement locally by Abies lasiocarpa, paralleling age and composition studies of modern forest stands in the region. This study is a proof of concept that, when applied to longer sedimentary records, could produce a detailed record of infestation for the Late Holocene or older time periods. This information will be useful to forest managers in efforts to plan for the effects of D. rufipennis infestations, and subsequent succession within high elevation conifer forests.     

Growth responses of three coexisting conifer species to climate across wide geographic and climate ranges in Yukon and British Columbia

This work aimed to compare radial growth–climate relationships among three coexisting coniferous tree species across a wide geographic and climate range from southern British Columbia (BC) to central Yukon, Canada. Tree-ring data were collected from 20 mature stands of white spruce (Picea glauca), lodgepole pine (Pinus contorta var. latifolia), and subalpine fir (Abies lasiocarpa). Linear relationships between annual growth variation and monthly and seasonal climate were quantified with correlation and regression analyses, and variation in climate–growth responses over a climatic gradient were quantified by regressing growth responses against local mean climatic conditions. Temperatures had more consistent and stronger correlations with growth for all three species than precipitation, but growth–climate responses varied among species and among sites. In particular, pine and fir populations showed different responses between BC and Yukon, whereas spruce showed a more consistent response across the study domain. Results indicate that (1) the response and sensitivity of trees to seasonal climate variables vary among species and sites and (2) winter temperatures prior to growth may have significant impacts on pine and fir growth at some sites. The capacity to adapt to climate change will likely vary among the study species and across climatic gradients, which will have implications for the future management of mixed-species forests in Yukon and BC.     

Climatic response of ring width and maximum latewood density of Larix sibirica in the Altay Mountains, reveals recent warming trends

? Introduction

Siberian larch (Larix sibirica) is a highly climate sensitive species. Presently, the Altay Mountains is covered by widespread forests dominated by Siberian larch and thus has a great potential for dendroclimatological studies. However, tree-ring network of the Altay Mountains has not yet been well developed. The development of the new chronologies and the knowledge about the influence of climatic variables on tree growth is needed.

? Method

X-ray densitometric techniques were applied to obtain ring width (RW) and maximum latewood density (MXD) of Siberian larch from two upper tree line sites in the Altay Mountains, China. Climatic responses in ring widths and maximum latewood densities from the Altay Mountains (China, Russia, and Mongolia) were investigated by simple correlation analyses. To assess the common growth forces among the individual sites of the Altay Mountains, simple correlation, principal component analyses, and spatial correlation analysis were applied over the common period of the chronologies.

? Results

Ring width and maximum latewood density increases with decreasing precipitation, increasing temperature from late spring to late summer during the growing season. Based on the results of principal component analyses and spatial correlation analysis, summer temperature (June?CJuly) is the most important forces on the Siberian larch growth of the Altay Mountains. The growth of Siberian larch in the Altay Mountains captures the current warming trend. The growth of Siberian larch did not clearly lose its sensitivity under most recent warming in our study areas.

? Conclusions

The new MXD chronologies is presently the longest, absolutely dated, tree-ring density record yet developed from China. The climate response analysis shows that the RW and MXD of Siberian larch have strong responses to temperature in the growing season. Thus, MXD and RW of Siberian larch provides the best information for climate reconstruction in the warm season. Tree-rings of Siberian larch allow detecting the recently observed warming trend and putting it into the long-term climatic context in the Altay Mountains, due to the strong growth sensitivity to temperature change.     

Effects of soil moisture and species composition on growth and productivity of trembling aspen and white spruce in planted mixtures: 5-year results

Aspen (Populus tremuloides Michx.) and white spruce (Picea glauca (Moench.) Voss) were planted 0.5 m apart in intimate mixtures in 5 × 4 m plots, with two moisture regimes—irrigation versus control—and five species compositions—pure aspen (Aw100), mixed aspen and spruce (Aw83Sw17, Aw50Sw50, Aw17Sw83), and pure spruce (Sw100), replicated six times. Fifth-year assessments indicated that irrigation increased individual tree growth (height, RCD, crown width), plot leaf area index (LAI), and wood biomass. Increased aspen composition reduced the availability of soil moisture and consequently the growth of individual trees. With increased aspen composition more growth was allocated to stem in aspen and to foliage in white spruce. Comparatively, aspen responded more to irrigation and thus their growth is more dependent on precipitation than that of spruce. Among the three growth variables assessed, height responded more to irrigation in both species. Equal mixtures and aspen-dominated mixtures in control plots had higher productivity in terms of total wood biomass in both absolute and relative terms. The implications of these findings are discussed in relation to managing aspen and white spruce mixedwood forests under increasing drought expected as a result of climate change.     

Evaluation of competitive effects of green alder,willow and other tall shrubs on white spruce and lodgepole pine in Northern Alberta

In boreal forests of western Canada, lodgepole pine (Pinus contorta Dougl. ex. Loud.) and white spruce (Picea glauca (Moench) Voss) often grow together with numerous tall shrubs such as green alder (Alnus crispa (Ait.) Pursh) and little-tree willow (Salix spp.). In an area south of Grande Prairie, Alberta, Canada, we examined the effects of shrubs, herbs and other trees on nutrient and light availability and growth of white spruce and lodgepole pine. For white spruce the best competition measure (tested against volume increments of the past 3 years) was visually estimated % ground cover times the height of the competitor (VCHT) with light (DIFN) ranking in third place. For lodgepole pine, DIFN was the best competition measure for predicting volume increment and the best competition index was again VCHT. Taller conifers had a stronger competitive effect than tall shrubs, with their effect on white spruce being larger than that on lodgepole pine.     

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.     

Relationships between climate and tree radial growth in interior British Columbia,Canada

Climate is a main driving factor of the formation of annual tree-rings, but which climatic variables are the most influential on radial growth may vary among species and sites. To explore these interactions, tree-ring chronologies along a major elevation gradient were examined for three tree species in southern interior British Columbia (Canada): Pseudotsuga menziesii, Pinus contorta, and Picea glauca × engelmannii. We used correlations and linear and multiple regressions to explore the relationships between tree-ring radial growth and climate variables in the area from 1922 to 1997. All correlation coefficients between ring chronologies and monthly climatic variables were medium to low (from −0.3 to 0.4); nevertheless, moderate but significant trends could be identified. Multivariate models explained up to 53%, 43% and 32% of radial growth variability for P. contorta, P. menziesii and P. glauca × engelmannii, respectively. All three species showed similar radial growth–climate patterns across the elevational gradient, but they had different details that made ring width–climate relationships species-specific. Precipitation-related variables were more related to radial growth at low-elevations, changing into temperature-related variables at high-elevations. Tree-ring width for all three species was primarily and significantly affected by climate variables from the year previous to the growing season and only secondly by current year conditions, but the critical months varied for different species and altitudes. Winter precipitation also affected radial growth, either as a source of water or as a possible agent of physical damage. Although our work showed significant climate influences on breast height tree radial growth, our results also indicated that other site factors such as microclimate or stand dynamics can be as or more important than climate variability.     

The spreading of the S type of Heterobasidion annosum from Norway spruce stumps to the subsequent tree stand

The occurrence of Heterobasidion annosum in stumps and growing trees was investigated on 15 forest sites in southern Finland where the previous tree stand had been Norway spruce (Picea abies) infected by H. annosum, and the present stand was either Scots pine (Pinus sylvestris), lodgepole pine (Pinus contorta), Siberian larch (Larix siberica), silver birch (Betula pendula) or Norway spruce 8–53 years old. Out of 712 spruce stumps investigated of the previous tree stand, 26.3% were infected by the S group and 0.3% by the P group of H. annosum. The fungus was alive and the fruit bodies were active even in stumps cut 46 years ago. In the subsequent stand, the proportion of trees with root rot increased in spruce stands and decreased in stands of other tree species. On average, one S type genet spreading from an old spruce stump had infected 3.0 trees in the following spruce stand, 0.5 trees in lodgepole pine, 0.3 trees in Siberian larch, 0.05 trees in Scots pine and 0.03 trees in silver birch stand. Although silver birch generally was highly resistant to the S type of H. annosum, infected trees were found on one site that was planted with birch of a very northern provenance.     

Sources of within- and between-stand variability in specific leaf area of three ecologically distinct conifer species

Specific leaf area (SLA) is an important ecophysiological variable, but its variability within and between stands has rarely been simultaneously examined and modeled across multiple species. Extensive datasets on SLA in coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco), hybrid spruce (Picea engelmannii Parry × Picea glauca (Moench) Voss × Picea sitchensis (Bong.) Carr.), and ponderosa pine (Pinus ponderosa Dougl. ex P. & C. Laws.) were used to estimate variability of SLA within a canopy and its relationship to tree- and stand-level covariates, and to predict SLA at various locations in tree crowns. Also, in the case of hybrid spruce, variation in SLA due to different relative horizontal lengths from the bole was examined. In all species, SLA systematically increased from tree tip to crown base and decreased with foliage age class. Cardinal direction did not have a highly significant influence in either Douglas-fir or hybrid spruce, but SLA did significantly decrease from branch tip to bole in hybrid spruce. Tree- and stand-level (e.g. density, site index) factors had relatively little influence on SLA, but stand age did have a significant positive influence. For ponderosa pine, a significant relationship between canopy mean current-year SLA and carbon isotope discrimination was also found, suggesting the importance of water stress in this species. An equation was fitted to estimate SLA at various points in the canopy for each species and foliage age class using absolute height in the canopy, relative vertical height in the tree, and stand age.