Why and where do adult trees die in a young secondary temperate forest? The role of neighbourhood

? The density and identity of tree neighbourhood is a key factor to explain tree mortality in forests, especially during the stem exclusion phase.

? To understand this process, we built a logistic model for mortality in a spatially explicit context, including tree and neighbourhood predictors. Additionally, we used this model to build mortality risk frequency distributions. Finally, we tested this model against a random mortality model to predict the spatial pattern of the forest.

? Annual mortality rate was high for pedunculate oak (Quercus robur, 6.99%), moderate for birch (Betula celtiberica, 2.19%) and Pyrenean oak (Q. pyrenaica, 1.58%) and low for beech (Fagus sylvatica, 0.26%). Mortality risk models for pedunculate oak and birch included stem diameter, tree height, canopy position and neighbourhood. Mortality was affected by the specific nature of the neighbourhood showing a clear competitive hierarchy: beech > pedunculate oak > birch. Models based on random mortality and logistic regression model were able to predict the spatial pattern of survivors although logistic regression predictions were more accurate.

? Our study highlights how simple models such as the random mortality one may obscure much more complex spatial interactions.

     

Lifespan and mortality of old oaks — combining empirical and modelling approaches to support their management in Southern Sweden

? Old oaks (Quercus robur L.) play an important role in the southern Scandinavian landscape by providing habitat for a wide range of species, a large proportion of them being currently on the National Redlists.

? To provide support for the management of these trees, we review data on oak mortality and formulate a mortality-driven stochastic model analysing interactions between mortality rate, oak recruitment rate into 100–150 age class, and amount of oaks older than 200 years.

? Empirical annual mortality rates varied between 0 and 13% with average 1.68%. Trees older 200 years had an average mortality rate of 1.1%. Oaks in the high density forests showed higher mortality (3.2%) as compared to the trees growing in the low density forests (1.2%). A 400-year long modelling exercises indicated that under current mortality rates (regular mortality being centred around 1% annually; and irregular mortality 7% with average return time of 13 years) the long-term maintenance of 20 trees older than 200 years per ha would require an input rate of 1 to 5 trees × year^?1 × ha^?1 into the 100–150 years old class.

? The modelling highlighted the importance of initial oak abundance affecting amount of old trees at the end of shorter (100 years) simulation period.

     

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.

     

Integration of water transport pathways in a maple tree: responses of sap flow to branch severing

? It has been known for a long time that sectored and integrated patterns of vascular systems exist in different species and even within the same tree, depending on its age and history. However, very few publications consider the topology of the vascular pathways between roots and branches.

? Some results on this important aspect of the vascular system are presented in this paper. They have been obtained with adult maple trees by directly studying the water movement in the stem and root xylem with the heat field deformation (HFD) method for sap flow measurements.

? Multi-point HFD sensors were installed at different heights of a Norway maple tree (Acer platanoides L.) along its stem axis. Single-point HFD sensors were installed in three small lateral roots of another sample maple. Experimental treatments (branch severing) triggered changes in sap movement in the stem and root sapwood.

? The sample trees belong to the group with an integrated transport system (“integrated pipes”), sharing stem space on both sides of the tree to supply two large parts of the crown with water from each root sector. Nevertheless, conducting pathways had their autonomy for axial transport and the pipe model theory describes the vascular system of the studied trees well. Thus, the integration of axial transport in the stem xylem should presumably occur through the cross-grained network of axial vessels.

     

Analysis of spatial patterns of oak decline in cork oak woodlands in Mediterranean conditions

? Cork oak mortality is a recurrent problem in southwestern Portugal. Despite the perception of increasingly visible damage in oak woodlands on drought-prone sites, the role of the various environmental factors in their decline is not clear.

? To describe the spatial patterns of cork oak (Quercus suber L.) mortality, a cork oak mortality index (MI) was determined for each landscape feature (agroforestry system, soil type, slope and aspect) using a GIS approach. To achieve this goal, a logistic regression model was formulated analyzing interactions between landscape attributes and allowing a prediction of cork oak mortality.

? Maximum values of MI were found in (i) shrublands and open woodlands with shrub encroachment (MI 6 and 3, respectively), where competition for soil water between tree and understory increases; and (ii) on lower slopes in the rounded hilltops and smooth hillsides or shallow soils where access to groundwater resources during summer drought is difficult.

? The model highlighted the importance of the agroforestry systems on cork oak mortality and may be used to identify sensitive areas where mitigation actions should be employed in a scenario of increasing drought severity in these Mediterranean ecosystems.

     

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.

     

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.

     

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.

     

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.

     

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.

     

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.

     

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.

     

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.

     

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.

     

Density dependence on tree survival in an old-growth temperate forest in northeastern China

? Density dependence is a major mechanism for shaping plant communities. However, its role in regulating diverse, mixed natural tree communities is less certain.

? In this study we investigated density-dependent effects in a large-scale (25 ha) old-growth temperate forest in northeastern China. Spatial patterns of neighborhood distribution in the plot were analyzed using various methods for inferring competition, including (1) pair correlation function to determine spatial patterns of pre-mortality and post-mortality and (2) neighborhood analysis of individuals to examine the extent to which tree survival is correlated with other covariates.

? Results showed that, for common species, 3 of 5 canopy species and 3 of 8 midstory and understory species were random in mortality. Negative density-dependent mortality was not found when trees reach 1 cm in DBH. There was no significant correlation for canopy species between tree survival and conspecific abundance, but largely positive correlations for midstory and understory species. In contrast, tree survival was found to negatively correlate with conspecific basal area for most species, indicating strong intraspecific competition. No strong interspecific density dependence was found in the forest.

     

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.

     

Influence of annual weather on growth of pedunculate oak in southern Sweden

? A network of oak (Quercus robur L.) chronologies containing 49 sites and 635 single trees was analysed to identify weather variables affecting annual tree-ring increment dynamics in southern Sweden during 1860–2000.

? We analysed (1) the growth response of oak to non-extreme weather, and (2) the temporal and spatial patterns of regional growth anomalies (pointer years) and associated climatic extremes resolved on a monthly scale.

? Growth was controlled by precipitation in the current (June–July) and the previous growing season (August) in 48% and 22% of all sites, respectively. Temperature during July of the current year and August of the previous year was negatively correlated with growth in 29% and 43% of the sites, respectively. Growth was positively correlated with temperature in October of the previous season in 72% of the sites. The most extensive growth anomaly occurred in 1965 and was probably caused by intrusion of cold Arctic air masses into the region at the end of March that year.

? During climatically non-extreme years, oak growth is driven mostly by the dynamics of summer precipitation. Many of the negative growth anomalies, however, were associated with temperature extremes. Southern Swedish oak pointer years tend not to coincide with the pan-European oak pointer years.

     

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.

     

Interactive responses of Quercus suber L. seedlings to light and mild water stress: effects on morphology and gas exchange traits

? The combined effect of water stress and light on seedlings of forest species is a key factor to determine the best silvicultural and afforestation practices in the Mediterranean area.

? The aims of this work was (1) to determine the optimal light level for the early development of cork oak seedlings under mild water stress and (2) to test if the combined effect of water stress and light followed the trade-off, the facilitation or the orthogonal hypothesis.

? Shade reduced instantaneous photosynthetic rates and water use efficiency in cork oak. However, seedlings grown under moderate shade (15% of full sunlight) were capable to accumulate similar amount of biomass than those grown under more illuminated environments by increasing their specific leaf area. Absolute differences in net photosynthesis between light treatments were higher in well watered than in water stressed seedlings. However, the impact of both factors on overall growth was orthogonal.

? We concluded that cork oak development is impaired under deep shade (5% of full sunlight) but it can be optimal under moderate shade (15% of full sunlight) even under moderate water stress. Implications of these patterns on regeneration, cultivation and afforestation of cork oak are discussed.