Using cover measurements to estimate aboveground understorey biomass in Maritime pine stands

? Understorey plays a major role in forest fluxes and stocks balances, however this compartment is generally poorly quantified. Our objectives were to establish models to estimate understorey biomass using vegetation cover measurements and to investigate upscaling methodologies from stand to regional level.

? Understorey aboveground biomass measurements were undertaken in Maritime pine stands of mesohygric, mesic and dry moorlands in South West France.

? Average biomass stock in this compartment was estimated to 3.50 t DM ha^?1. The more abundant species groups varied with moorland types, with a higher relative contribution of herbaceous species (23.3%), bracken (59.2%) and mosses (31.6%) for mesohygric, mesic and dry moorlands, respectively. For each species group, we established significant relationships to estimate biomass using a volumetric index, based on cover and height measurements. No relationship between stand characteristics and understorey biomass was founded. We investigated the upscaling of these estimations to a several thousands hectare area using understorey cover measurements done along a regular spatial grid. The only significant correlation linked one satellite vegetation index to understorey biomass.

? We successfully developed empirical relationships to estimate the understorey biomass at the stand level. Further investigations could focus on the analysis of understorey variability over a finer space grid and the potential use of satellite vegetation indexes.

     

The potential of plantations of Terminalia superba Engl. & Diels for wood and biomass production (Mayombe Forest, Democratic Republic of Congo)

? In the 1940s–1950s, large limba (Terminalia superba Engl. & Diels) plantations were established in the Democratic Republic of Congo to reduce the pressure on the natural forests.

? The objective of this study was to evaluate the potential of these long-rotation plantations as production forests (timber) and carbon sinks.

? Five different plantations, between 50 and 58 years old, were sampled. Over a sample surface of more than 73 ha, the diameter above buttresses of 2 680 trees, bole height of 265 trees and tree height of 128 trees was measured.

? To estimate the commercial volume, a nonlinear power law regression was used (R ² = 0.95). A power law variance function was applied to counter heteroscedasticity of the residual plot. Estimates of commercial tree and stand volume at 50 to 58 y were 5.6 ± 4.1 m³ and 183.9 ± 135.0 m³ ha^?1. Stand volumes appear low but are explained by a large decrease in tree density. However, the mean volume increment of 3.2–3.7 m³ ha^?1 y^?1 corresponds well with teak plantations of a similar age. For limba, aboveground biomass and carbon estimates of this study (resp. 108.4 and 54.2 Mg ha^?1) differ significantly from those of existing aboveground biomass models (resp. 135.7–143.9 Mg ha^?1 biomass and 67.9–72.0 Mg ha^?1 C). All aboveground biomass and carbon estimates for T. superba stands were lower than for the estimates of young fast-growing plantations like Tectona grandis L. f., Eucalyptus spp. and Acacia spp. (≤ 30 y).

     

Effects of some ecological variables on carabid communities in native and non native forests in the Ibaizabal basin (Basque Country: Spain)

? Carabid beetles were investigated at five different forest types in the Ibaizabal basin (northern Spain). The landscape is characterized by the presence of remnants of native forest surrounded by conifer plantations.

? Carabids were trapped in 52 stands of mixed forest, beech forest, holm oak forest, mixed pine and Monterey pine plantations in 2005 and 2006. The main objectives of the study were: compare carabid diversity, recognise the characteristic species, and study the effects of ecological variables on carabid assemblages in the different forest types.

? No significative differences in species abundance, richness and diversity were found among the studied forests. Most of the trapped beetles were identified as forest generalists, nevertheless some native and non-native forest specialist species were also found. Distribution of carabid communities overlapped and, except for beech forest, no specific communities were distinguished. Altitude, percentage of grass coverage and temperature were the main variables influencing species distribution.

? The results suggest high habitat homogeneity, caused by regeneration in pine plantations of the indigenous understorey, and by poor habitat quality in native forest (patchy remnants enclosed in conifer plantations). This situation could explain the similar carabid diversity.

     

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.

     

Tree diameter, height and stocking in even-aged forests

? Empirical observations suggest that in pure even-aged forests, the mean diameter of forest trees (D, diameter at breast height, 1.3 m above ground) tends to remain a constant proportion of stand height (H, average height of the largest trees in a stand) divided by the logarithm of stand density (N, number of trees per hectare): D = β(H ? 1.3)/ ln(N).

? Thinning causes a relatively small and temporary change in the slope β, the magnitude and duration of which depends on the nature of the thinning.

? This relationship may provide a robust predictor of growth in situations where scarce data and resources preclude more sophisticated modelling approaches.

     

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.

     

Properties of boundary-line release criteria in North American tree species

? Boundary line release criteria are increasingly applied to evaluate forest disturbance histories from tree-ring data. However, a number of important properties central to the technique have not been evaluated, including: (i) the ability of boundary line release criteria to standardize releases across various sites, species, and tree life stages (ii) the minimum sample sizes necessary for developing boundary lines, and (iii) the degree to which the criteria can resolve the degree of crown exposure following a disturbance event.

? In an analysis of eleven North American tree species, boundary line release criteria do not fully compensate for declines in release response a tree experiences with increasing age and size, with the exception Tsuga canadensis.

? A bootstrapping analysis indicates that approximately 50 000 ring width measurements are necessary to develop boundary line release criteria for a given species.

? In a Quercus prinus stand, boundary line release criteria better predict the degree of crown exposure following a disturbance than an earlier running mean technique.

? Despite certain limitations, boundary line release criteria have the potential to standardize release calculation across most life stages of a tree, and possibly among sites and species.

     

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.

     

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.

     

Epicormic sprout development in pruned coast redwood: pruning severity, genotype, and sprouting characteristics

? Young coast redwood (Sequoia sempervirens (D. Don.) Endl.) trees were pruned to various heights to examine the effect of pruning severity on epicormic sprouting. Seven separate stands were used with as many as six treatments per stand in coastal Humboldt County, California, USA.

? Epicormic sprout development was affected by pruning severity but primarily at the most severe pruning treatments that removed all but the branches in the top 15% of tree height. Less severe treatments produced sprouts but the number and size of these sprouts were comparable to unpruned trees.

? Natural clonal patterns were also used to explore patterns of sprouting between genotypes. Linear mixed-effects models were developed to predict sprouting frequency as a function of pruning severity while accounting for the nested data structure (i.e., stem sections sampled nested within genotypes within treatments within sites).

? Comparing variances attributed to each of these random effects indicated that at any level of pruning severity, differences in epicormic sprouting between genotypes and sites expressed soon after pruning had disappeared after six growing seasons. Epicormic branches were more common two years after pruning than six years indicating many branches were dying. Branches were more common in the middle of the pruned bole, possibly because of competition from basal sprouts and the expanding tree crown.

     

Effects of stand density on ecosystem properties of subalpine forests in the southern Rocky Mountains, USA

? Mixed coniferous, subalpine forest communities in the Rocky Mountains are historically dense and have experienced infrequent, high-severity fire. However, many of these high-elevation stands are thinned for a number of perceived benefits.

? We explored the effects of forest stand density on ecosystem properties in subalpine forests in Colorado, USA, 17–18 y after forests were managed for timber.

? Forest structure significantly altered the composition and chemical signature of plant communities. Previously managed stands contained lower density of overstory trees and higher ground cover compared to paired reference stands. Foliar phenolic concentration of several species was negatively related to basal area of overstory trees. Furthermore, reductions in stand density increased total foliar phenolic:nitrogen ratios in some species, suggesting that gap formation may drive long-term changes in litter quality. Despite significant changes in forest structure, reductions in stand density did not leave a strong legacy in surface soil properties, likely due to the integrity of soil organic matter reserves.

? Changes in forest structure associated with past management has left a long-term impact on plant communities but has only subtly altered soil nutrient cycling, possibly due to trade offs between litter decomposability and microclimate associated with reductions in canopy cover.

     

Preliminary study on the assessment of deadwood volume by the French national forest inventory

? As the French national forest inventory does not currently monitor the lying compartment of the forest deadwood, the figures obtained on this topic are therefore partial. This study provides cost estimates and guidelines for assessing stumps, and standing and lying deadwood.

? Comparisons were made between a fixed-area sampling method and a line-intersect one. LIS was judged more time-efficient, especially in dense understorey. Computer simulations were performed in order to estimate the gain in precision with increasing transect lengths. The results showed a continuous improvement in precision associated with increases in transect length. The longest transect tested (50 m) still had a large coefficient of variation, suggesting that improvement in precision at the plot level could still be gained with longer transects.

? Therefore, from a practical standpoint in terms of fieldwork, we suggest that on a national scale lying deadwood should be measured by line-intersect sampling, whereas stumps, standing dead trees and snags can be monitored using standard fixed-area plots. To meet needs at the national level, we consider that local imprecision could be compensated for by the large number of plots measured each year.

     

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.

     

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.

     

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.

     

Potential use of pine plantations to restore native forests in a highly fragmented river basin

? In forests, the substitution of broadleaf species by conifers can reduce biodiversity because coniferous forests generally provide less diverse vascular understories than broadleaf forests. However, in some cases, former pine plantations might be useful for restoring native forests. We compared plant species composition on the plot scale in natural beech and mixed oak forests with that in plantations of Pinus radiata. Links between plant diversity and landscape parameters (patch size, fractal dimension and distance to the nearest patch of the same type) were investigated.

? The objective of this study was to evaluate the use of pine plantations for restoring native diversity in a zone where native forests are very fragmented.

? Similar to oak forests, plant diversity in pine plantations was high, mainly due to the presence of generalist species. Some species characteristic of oak forests also appeared in pine plantations, suggesting the onset of natural forest regeneration.

? These results suggest that pine plantations could be used to promote natural regeneration of original oak forests. Moreover, residual native stands should be conserved as important sources of native species and their seeds.

     

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.

     

Climate-growth variability in Quercus ilex L. west Iberian open woodlands of different stand density

? We present the longest tree-ring chronology (141 y) of Quercus ilex L. (holm oak), and discuss the species climate-growth relationships and the influence of stand density on tree sensitivity to climate.

? Similarly to Quercus suber L., the most influential climatic variables upon holm oak growth were late spring and early summer precipitation, which enhanced growth, and high temperatures in the previous August and current July, which negatively affected growth.

? High density stands responded to similar climatic factors as low density stands, but their response was generally weaker. Holm oak sensitivity to climate has increased in recent decades, which might be related to increasing temperatures in the region. Sensitivity was higher in low density stands. Additionally, the effect of summer stress on growth seems to have increased during the same period, similarly to other species in the Iberian Peninsula, suggesting that trees are more vulnerable to climatic changes.

? Stand density could buffer the response to climate by smoothing climatic extremes. Nevertheless, the effect of competition might reverse this positive effect at the individual tree level. Precautions should be taken before providing management guidelines regarding the effect of climate change and stand density on holm oak.

     

Influence of several tree traits on rainfall partitioning in temperate and boreal forests: a review

? In forests, rainfall is partitioned into intercepted water (IW), throughfall (TF) and stemflow (SF). We reviewed the majority of published works comparing water flows among tree species in temperate and boreal forests to test the effect of several tree traits on water flows.

? We hypothesized that water flows differed between evergreen and deciduous species, and according to successional status and bark roughness. We also investigated that water flows would be explained by stand variables such as basal area.

? Linear mixed models fitted on reviewed data showed that evergreens had a lower TF than deciduous trees (decrease of 13.9% of total precipitation year-round and 8.4% over the growing period). Similar results were found for conifers compared to broadleaves. TF also declined with the successional status from pioneer to late-successional tree species. SF decreased with bark roughness from smoother to rougher bark. Evergreens had water flows that were dependent on age of the stand, especially for TF which increased by 15.6% of total precipitation from young to adult forests.

? The large scale of TF differences between tree genera together with specific transpiration amounts and rooting features highlighted in other studies should result in significant differences in soil water content among tree species. This may have consequences on stand fitness and growth, and understory vegetation.

     

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.