Effects of climate variables on intra-annual stem radial increment in Pinus cembra (L.) along the alpine treeline ecotone

? Within the alpine treeline ecotone tree growth is increasingly restricted by extreme climate conditions. Although intra-annual stem growth recorded by dendrometers can be linked to climate, stem diameter increments in slow-growing subalpine trees are masked by changes in tree water status.

? We tested the hypothesis that intra-annual radial stem growth in Pinus cembra is influenced by different climate variables along the treeline ecotone in the Austrian Alps. Dendrometer traces were compared with dynamics of xylem cell development to date onset of cambial activity and radial stem growth in spring.

? Daily fluctuations in stem radius reflected changes in tree water status throughout the treeline ecotone. Extracted daily radial increments were significantly correlated with air temperature at the timberline and treeline only, where budburst, cambial activity and enlargement of first tracheids also occurred quite similarly. A close relationship was detected between radial increment and number of enlarging tracheids throughout the treeline ecotone.

? We conclude that (i) the relationship between climate and radial stem growth within the treeline ecotone is dependent on a close coupling to atmospheric climate conditions and (ii) initiation of cambial activity and radial growth in spring can be distinguished from stem re-hydration by histological analysis.

     

Diminished vessel diameter as a possible factor in the decline of European ash (Fraxinus excelsior L.)

? The aim of this work was to examine the anatomy and functioning of secondary xylem in stems in relation to the decline of European ash (Fraxinus excelsior L.).

? We tested the hypothesis that declining trees show changes in the structure of wood, which result in impaired water transport.

? Anatomical analyses were carried out on wood samples (comprising all annual rings formed during the 30 years life of the analysed trees) collected at breast height from the main stem of healthy, weakened and dead ash trees. The width of annual wood increments, the diameter and density of earlywood vessels were measured and the theoretical hydraulic conductivity index through the secondary xylem calculated by application of Hagen-Poisseuille’s formula.

? Anatomical characteristics changed both with the age of trees and in response to unfavorable factors. The largest vessels were observed in healthy trees, which implied that they had the highest hydraulic conductivity index, whereas trees considered to be in decline produce smaller vessels and hence had reduced conductivity.

     

Tree mortality in a mixed deciduous forest in Northwestern Russia over 22 years

? We studied mortality rates of birch, small-leaved linden, pedunculate oak, Norway maple, black alder, common ash and Norway spruce, assessed through two inventories of 12 664 trees in the unmanaged mixed deciduous forests of Oranienbaum Park, northwestern Russia, in 1981 and 2003.

? Our objectives were to evaluate if (a) long-term mortality rates vary among species, and if (b) crown condition, age and tree size affect the probability of mortality.

? Over this period, tree mortality rates in the park varied annually from 1 to 3% for healthy and moderately healthy trees, and from 3.9 to 9.3% for declining trees. The lowest mortality rates were observed for small-leaved linden and oak (0.8 and 1.0%, respectively), and the highest rate for birch (2.7%). We found (1) a significant and consistent association between tree condition and the probability of mortality, and (2) significantly higher mortality rates in smaller trees of ash, maple and oak.

? The effect of species-specific biology on mortality rates in the Oranienbaum Park was largely overridden by the “health status” and environmental conditions of the trees (e.g. degree of crown shading). The observed rates were similar to the estimates from natural temperate deciduous forests in both Europe and North America, indicating similar intensity of mortality in these ecosystems under natural conditions.

     

Variations in water use by a mature mangrove of Avicennia germinans, French Guiana

? In the tropical intertidal zones, little is known on water uptake by mangroves. Transpiration rates are generally measured at leaf level, but few studies exist on water use at tree or stand levels.

? The objective of this study was to measure sap flow in trees of different sizes to appreciate the range of variation in water use that may exist in a site dominated by 80% mature Avicennia germinans.

? The results showed that from the dry to the wet season the mean water use increased from 3.2 to 5.3 dm³ d^?1 in small trees (DBH ～ 13 cm), from 11.5 to 30.8 dm³ d^?1 in medium trees (～24 cm) and from 40.8 to 64.1 dm³ d^?1 in large ones (～45 cm).

? Sapwood remained active up to a depth of 8 cm with radial variations within the stem. Weak correlations were obtained with VPD and net radiation.

? This study confirmed that transpiration was larger under low levels of salinity. Water use at stand level (～1900 living stems ha^?1) was estimated to be in the range of 5.8 to 11.8 m³ ha?1 d^?1 according to the season.

     

Gender, neighboring competition and habitat effects on the stem growth in dioecious Fraxinus mandshurica trees in a northern temperate forest

? This study analyses the sex ratios, and the effect of gender, neighboring competition and habitat factors on the stem growth of dioecious F. mandshurica trees in the Changbai Mountains of North-Eastern China.

? The general sex ratio of the flowering trees did not significantly deviate from 1:1. The sex ratio varied among age classes. A female-biased sex ratio was found in the age classes of 65–75 years and 85–95 years, but departure from 1:1 was not significant in other age classes.

? An analysis of variance revealed that the growth rate was extremely sensitive to gender and tree size, as shown by a highly significant gender-related and size-dependent effect. Male trees had a faster stem growth rate than female trees, and the assumption is that trade-offs between reproductive and vegetative processes can induce the difference in stem growth rate between genders.

? Stem growth rates are positively affected by soil moisture and tree size for both genders at all considered neighborhood distances. The growth rates of female trees were negatively affected by neighboring competition from other female trees at all neighborhood distances between 1 and 10 m, but were not significantly affected by male trees. The growth rates of male trees were negatively affected by neighboring competition from male trees for neighborhood distances between 3 and 10 m, but were not significantly affected by female trees.

? These results suggest that intrasexual competition (male-male competition and female-female competition) may cause difference in stem growth for both genders. Intersexual differences in sensitivity to neighboring plants are considered to be an important factor driving gender-specific growth patterns.

     

Extending sapwood — Leaf area relationships from stems to roots in Coast Douglas-fir

? Studies of allometric relationships between leaf area and the cross-sectional area (CSA) of sapwood in the stem have shed light on the structural and functional relationships between water-conducting and photosynthetic tissues.

? The purpose of this study was to test whether sapwood-leaf area relationships could be extended from stems to roots in coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Twelve trees were felled, their stumps were excavated, and the CSA of sapwood and heartwood were estimated for individual roots, entire root systems, and stem section.

? Root sapwood CSA was greater than sapwood CSA throughout the stem, and the ratio of leaf area to sapwood CSA (A _l :A _s ) was accordingly lower for root sapwood. The relationship between sapwood CSA and leaf area was more variable in roots and at groundline compared to crown base. Root A _l :A _s decreased with relative tree height (tree height/mean stand height).

? The strong allometric relationship between leaf area and the CSA of sapwood in the stem generally holds when extended to roots. The greater CSA of sapwood in roots versus stems may reflect differences in their roles in supporting the tree.

     

Stimulation of tree defenses by Ophiostomatoid fungi can explain attack success of bark beetles on conifers

? Our aim is to present why the hypothesis, that Ophiostomatoid fungi play an important role in the establishment of most bark beetle species on living conifers, is valuable.

? After summarizing knowledge about the relationships of bark beetles with conifers and fungi, we conclude that controversy results from misinterpretations when using fungal pathogenicity to demonstrate the role of Ophiostomatoid fungi in beetle establishment on host trees.

? We demonstrate that fungal pathogenicity is not the right parameter to appreciate the role of fungus in beetle establishment on host trees. We argue that artificial low density inoculations that allow the appreciation of fungus ability to stimulate tree defenses and thus to help beetles in overcoming tree resistance must be used in complement to mass inoculations. In both cases, results must be expressed in terms of tree defense stimulation rather than in terms of tree killing.

1. Fungal species stimulating tree defenses are generally not those that grow the best in the sapwood.
2. We argue that beetle development in the phloem, fungal invasion of the sapwood and phloem, and tree death, occur after tree defenses are exhausted, and that any fungus present in the beetle gallery could thus potentially invade the sapwood after defense exhaustion.

? We conclude that stimulation of the tree defense reactions in both the phloem and the superficial sapwood is a real benefit brought by fungi to the beetles during the first phase of establishment (overcoming tree resistance).

? Considering the origin of the bark beetle fungus associations attacking living trees and their general functioning based on stimulation of tree defenses, we develop three hypotheses:

1. any beetle species would be helped in its establishment in a given tree species by developing an association, even loosely, with a fungus species belonging to the Ophiostomatoid flora of that tree species;
2. the necessity of a considerably low level of tree resistance for fungus extension into the tree is the selection pressure that has led fungi to develop their intrinsic ability to stimulate tree defenses, through their ability to grow into the phloem. This association can be completed by antagonistic fungal species controlling extension of the previous fungal species in the tree tissues;
3. Beetle species using the strategy of overcoming tree resistance are associated with a fungal complex, of which species could assume three roles regarding relationships between beetles and trees: 1- to stimulate tree defenses in the phloem and superficial sapwood, 2- to grow into the sapwood after tree resistance is overcome, and 3- to control phloem extension of the first other two categories. Bringing nutrients to the beetle progeny can be a fourth role.

? We propose that bark beetle — Ophiostomatoid associations can be categorized, based on associations’ frequency and complexity while taking into account beetle aggressiveness. We show that a close correspondence exists between beetles’ aggressiveness and the ability of their main associated fungal species to stimulate the defenses of their host tree.

? We conclude with suggesting that most sapwood invading fungi might be “cheaters” which have taken advantage of the efficiency of the relationship between beetles and fungi that stimulate tree defenses.

     

Diachronic analysis of individual-tree mortality in a Norway spruce stand in the eastern Italian Alps

? Understanding tree mortality processes across time requires long term studies. Spatiotemporal patterns of mortality in a 200 years-old mono-layered Norway spruce stand were evaluated to determine what factors affected individual-tree mortality.

? We performed an analysis on two surveys (1993 and 2005) in a 1-ha permanent plot in the Paneveggio forest (Eastern Italian Alps). Tree diameter and age distribution between surveys were compared. We examined spatial patterns of living and dead trees before 1993, in 1993 and in 2005 using univariate and bivariate Ripley’s K(d) function, and a kernel estimator of local crowding. A logistic model was used to assess the effects of diameter, age, recent growth and competitive pressure on tree mortality.

? Spatial pattern analysis indicated mortality was associated to tree neighbourhood (neighbour effect at 2–5 m). An increment of regularization of tree spatial pattern occurred due to density-dependent mortality. Logistic regression showed tree diameter and recent growth were determinant on mortality risk during the monitoring period.

? Even if the stand is relatively aged, mortality dynamics are those typical of stem exclusion stage. Mortality was related to competitive dynamics, and small suppressed trees with slow growth rate had higher probability to die.

     

Is forest mushroom productivity driven by tree growth? Results from a thinning experiment

? Most of the edible forest mushrooms are mycorrhizal and depend on carbohydrates produced by the associated trees. Fruiting patterns of these fungi are not yet fully understood since climatic factors alone do not completely explain mushroom occurrence.

? The objective of this study was to retrospectively find out if changing tree growth following an increment thinning has influenced the diversity patterns and productivity of associated forest mushrooms in the fungus reserve La Chanéaz, Switzerland.

? The results reveal a clear temporal relationship between the thinning, the growth reaction of trees and the reaction of the fungal community, especially for the ectomycorrhizal species. The tree-ring width of the formerly suppressed beech trees and the fruit body number increased after thinning, leading to a significantly positive correlation between fruit body numbers and tree-ring width.

? Fruit body production was influenced by previous annual tree growth, the best accordance was found between fruit body production and the tree-ring width two years previously.

? The results support the hypothesis that ectomycorrhizal fruit body production must be linked with the growth of the associated host trees. Moreover, the findings indicate the importance of including mycorrhizal fungi as important players when discussing a tree as a carbon source or sink.

     

Which factors most influence heartwood distribution and radial growth in plantation teak?

? To investigate the effect of climate on radial growth in young plantation grown teak (Tectona grandis L.), growth ring width was measured in 105 trees and correlated to precipitation and temperature data.

? The social status of trees within the stand was also determined and cross-sectional area (CSA) for the trunk correlated to the proportion of heartwood (HW) within the tree. HW develops asymmetrically in leaning stems of some conifer species, but it is not known if this phenomenon also occurs in broadleaf species. Therefore, we measured HW proportion in leaning and straight stems, along with the number of growth rings in the HW.

? Annual ring width depended strongly on mean monthly temperature during the rainy season and the most significant relationships were found corresponding to the months of June and July. With regard to the weaker relationship between precipitation and radial growth, correlations were highest during the period of bud-break at the beginning of the rainy season.

? The very high stand density affected radial growth, particularly in suppressed trees, which responded little to thinning operations. HW formation was greatest in dominant trees, and was highly regressed with stem CSA.

? Therefore, rapid growth of young stands should be encouraged by reducing stand density. Asymmetric HW formation occurred in both leaning and straight trees, and was significantly greater along the upper sides of leaning stems. It is probable that this eccentric HW formation is linked to mechanical loading on the tree.

     

Physical properties of wood in thinned Scots pines (Pinus sylvestris L.) from plantations in northern Spain

? The physical properties of wood and the associated variations within and between trees were evaluated by analysing 770 small specimens of clear wood from 11 Pinus sylvestris L. trees thinned from 3 plantations.

? Within-tree variations in basic density or volumetric shrinkage increased with cambial age and decreased with increasing ring width. The effect of the height in the stem on wood properties was considered indirect and height was not included as an explanatory variable in the mixed models proposed to estimate basic density and volumetric shrinkage.

? The models had random components for the intercept parameter and explained 52.5% of the total variance in basic density and 56% of the total variance in volumetric shrinkage. Linear shrinkage in the direction of the grain was extremely variable.

? Between-tree variation and between-plot variation in the physical properties of wood were high, considering that all trees sampled were growing in similar sites and stands. It would then be desirable to predict physical properties of wood on living trees in order to use the quality of wood as a criterion for timber tree selection in thinnings.

     

Improving models of forest nutrient export with equations that predict the nutrient concentration of tree compartments

? The objective of this study was to explore the distribution of major nutrients (N, P, K, Ca and Mg) in the aboveground compartments of an intensively managed tree species (Pinus pinaster Ait.). A total of 53 trees were cut down in even-aged stands respectively 8, 16, 26, 32 and 40 years old. The nutrient concentrations of the aboveground compartments were analysed.

? Nutrient concentrations of foliage did not vary with any of the variables used, except needle age. Nutrient concentrations of living branches, stem bark, stem sapwood, stem heartwood, stemwood and stem decreased with increasing branch diameter, bark thickness, sapwood thickness and heartwood thickness, respectively. Beyond a certain value of the predictive variable (stem diameter ≈ 15 cm; branch diameter ≈ 2.5 cm), the concentration of all the nutrients stabilised.

? A 50 year-old pine stand was used to obtain a validation dataset for nitrogen concentration. For this nutrient, the regression relationships gave satisfactory estimates for most compartments (mean error = 12–25%) and particularly for the stem.

? A procedure is proposed to estimate the nutrient exports associated with harvests of Pinus pinaster biomass.

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Towards standardised crown condition assessment in poplar plantations

? This work aims at developing new tools for rapid assessment of forest health indicators in poplar plantations.

? Crown transparency and discoloration were visually evaluated in all trees of four 15 m-radius sub-plots in 32 poplar clonal plantations, which were chosen according to a factorial scheme with three factors: tree age, site quality and understorey vegetation management. A subset of trees was assessed using digital photos processed with a semi-automatic image analysis system (the CROCO software) in order to compare visual and digital crown transparency estimates.

? Poplar crown conditions were better in young stands and rich sites. Harrowing understorey vegetation improved tree health in poor sites. Samples of 20 trees per stand provided the same information about crown transparency and discoloration as 60 trees. Calibration curves of digital crown transparency estimates were successfully fitted against visual crown transparency estimates. The same effects of stand age and site quality could be detected with digital crown transparency as response variable.

? The use of digital photos processed with CROCO in ca. twenty trees per stand is therefore recommended to accurately and objectively monitor crown condition in clonal poplar plantations.

     

Diversity of deciduousness and phenological traits of key Indian dry tropical forest trees

? In seasonally dry tropical forests deciduousness (leaflessness) is an important strategy of trees to survive in water stress period during summer. Deciduousness is a reflection of interacted effect of seasonal drought, tree characteristics and soil moisture conditions.

? The present study aims to document the diversity in leaf pheno-phases in terms of duration of deciduousness (which is reciprocal to growing season length), wood density, leaf mass per area (LMA) and leaf strategy index in 24 important tree species growing in the Vindhyan dry tropical forest in India.

? On the basis of phenological observations, the tree species were categorized into two main groups: leaf exchanging species exhibiting overlapping periods of leaf fall and leaf flush, and deciduous species whose timings of leaf flush and leaf fall differ resulting in a time lag (deciduousness) between the completion of leaf fall and initiation of leaf flush. Presence of wide range of deciduousness duration (from ca. a week to 7 months) among dry tropical trees indicates large variations in their growing season length. In the tree species studied, as the duration of deciduousness increased, leaf flushing period decreased significantly but leaf fall period showed little variation.

? Differing deciduousness in tree species exhibited substantial differences in their leafing (vegetative growth) pattern, as reflected by ratio of durations of leaf flush to leaf fall (leaf strategy index). Across different species, duration of deciduousness was significantly positively correlated with leaf strategy index, and significantly negatively correlated with both wood density and LMA.

? Wide variations in deciduousness, leaf strategy index, wood density and LMA in the 24 species investigated indicate considerable functional diversity in tree species growing in Vindhyan dry tropical region. Variation in seasonal duration of deciduousness among species is reflections of differences in tree functional traits like stem wood density, leaf strategy index and LMA.

     

Black pine (Pinus nigra Arn.) growth divergence along a latitudinal gradient in Western Mediterranean mountains

? Most studies of tree-growth and climate report positive responses to global warming in high latitudes and negative responses at lower ones.

? We analyzed tree-ring width of Pinus nigra Arn. along a 500 km latitudinal transect in the Iberian Peninsula to study the temporal trend and climate forcing in tree radial growth during the last century.

? Tree growth was enhanced by cool summers and moist cold seasons. Increased moisture stress has decreased tree growth rates. However, we present evidence of growth increases in some trees in all sampled populations after 1980’s. Climate change negatively (positively) affected between 72% (5%) of trees in the southern populations and 40% (25%) in the north Trees with positive growth trends were favored by winter temperatures and their abundance was inversely correlated with forest productivity.

? Our findings add evidences of tree growth divergence in the Mediterranean basin and show the gradual transition between forests where positive (temperate and boreal) and negative (Mediterranean) growth trends dominate.

     

Effects of the lateral growth rate on wood quality of Gmelina arborea from 3.5-, 7- and 12-year-old plantations

? Awareness of the shortage of fossil resources leads to an increasing demand for woody biomass. We investigated the feasibility of using fast-growing Gmelina arborea wood for material production. Gmelina arborea wood samples were collected from trees of varying cambium ages in Indonesia, from 3.5-, 7- and 12-year-old plantations.

? The lateral growth rate and the cambium age did not significantly affect the longitudinal released strain of the growth stress, xylem density, or microfibril angle at the outermost surface of the secondary xylem at any sampling site. However, fiber length in the 3.5-year-old plantation tended to be shorter in smaller diameter trees, whereas in larger diameter trees it was almost the same as that in trees from the 7- and 12-year-old plantations. This suggests that smaller diameter trees in the 3.5-year-old plantation had not yet produced mature wood.

? Xylem qualities had already reached values appropriate for harvesting, except in the smaller diameter trees from the 3.5-year-old plantation. This indicates that the larger diameter trees had already matured, regardless of their cambium age. These results suggest that the next step is to develop silvicultural treatments to increase the lateral growth rate during the early growing stage, in order to produce as much mature wood as possible, as quickly as possible.

     

Reconstruction of soil pH by dendrochemistry of Masson pine at two forested sites in the Pearl River Delta, South China

? Soil acidification is a global concern. Base cation (Ca, Mg, Mn and Al) concentrations, as well as their molar ratios, were measured in tree rings of Masson pines (Pinus massoniana L.) from Dinghushan and Xiqiaoshan to reconstruct the historical changes in soil acidification in the Pearl River Delta, south China.

? In Dinghushan, Ca and Mn xylem concentrations yielded significant correlations with the higher soil pHs, and Ca and Mg with the lower soil pHs. In Xiqiaoshan, only xylem Mn concentration correlated significantly with the soil pH in both layers. Significant correlations between xylem molar ratios and soil pH were revealed for Ca/Mn, Mg/Mn, Mn/Al and Ca/Al in the upper soils and for the same ratios except for Ca/Al in the deeper soils in Dinghushan. The same correlations were revealed for Ca/Mn, Mg/Mn and Mn/Al at both depths in Xiqiaoshan.

? Soil pHs at 0–10 cm and 10–40 cm depths at the study sites were reconstructed by the molar ratios of Mg/Mn and Ca/Mn in the xylem of Masson pines, respectively.

? Cation molar ratios in the xylem of Masson pine were superior to the single Ca, Mg, Mn and Al for soil acidity reconstruction in the Delta of China.

     

A comparison of mortality rates between top height trees and average site trees

? Although comparisons between methods of selecting trees for site index estimates are well documented in the literature, little is known on mortality rates of different canopy tree cohorts used for that purpose.

? This study was initiated to test the hypothesis that the mortality rates of top height trees are lower than those of codominants only or a combination of codominant and dominant trees. To test this hypothesis, we used records from a network of permanent sample plots in Québec and studied the fate of different cohorts of site trees for five different species.

? Our results did not show clear evidence of lower mortality rates for top height trees. Instead we found that depending on the species, top height trees have lower (Populus tremuloides, Pinus banksiana), higher (Picea mariana, Abies balsamea) or equal mortality rates (Betula papyrifera) than codominant trees or codominant and dominant trees combined.

? These results suggest a tendency for shade intolerant species to maintain lower top height tree mortality rates over time when compared to shade tolerant species. In the latter case, it is also shown that spruce budworm epidemics (Choristoneura fumiferana) did not change the pattern of mortality rates of site trees of A. balsamea.

     

Comparison of model forms for estimating stem taper and volume in the primary conifer species of the North American Acadian Region

? The performance of ten commonly used taper equations for predicting both stem form and volume in balsam fir [Abies balsamea (L.) Mill], red spruce[Picea rubens (Sarg.)], and white pine[Pinus strobus (L.)] in the Acadian Region of North America was investigated.

? Results show that the Kozak (2004) and Bi (2000) equations were superior to the other equations in predicting diameter inside bark for red spruce and white pine, while the Valentine and Gregoire (2001) equation performed slightly better for balsam fir.

? For stem volume, the Clark et al. (1991) equation provided the best predictions across all species when upper stem diameter measurements were available, while the Kozak (2004) and compatible taper equation of Fang et al. (2000) performed well when those measurements were unavailable.

? The incorporation of crown variables substantially improved stem volume predictions (mean absolute bias reduction of 7–15%; root mean square error reduction of 10–15%) for all three species, but had little impact on stem form predictions.

? The best taper equation reduced the predicted root mean square error by 16, 39, and 45% compared to estimates from the widely used Honer (1965) regional stem volume equations for balsam fir, red spruce, and white pine, respectively.

? When multiple taper equations exist for a certain species, the use of the geometric mean of all predictions is an attractive alternative to selecting the “best” equation.