Increasing Tree Size Research Articles

Maximum density patterns in two natural forests: An analysis based on large observational field studies in China

Previous studies on self-thinning have almost exclusively dealt with even-aged monocultures and the aim of such studies was to present some limiting relationship between average plant size and density. Our knowledge regarding density patterns and maximum densities in uneven-aged natural forests, and the ecological mechanisms that produce such patterns, is still very limited. This study analyses the density patterns in two species-rich forests in North-Eastern China. Each of the two 28 hectare field plots with mapped trees was subdivided into (320 and 312) cells, each cell measuring 30m×30m. The limiting relationships for the two study areas were estimated using exponential models and percentile regression. The allometric exponents between tree size and tree density are surprisingly similar for Jiaohe (−1.17) and Liangshui (−1.14), and considerably lower than those found in even-aged plantations. Another objective of the study was to study the relationship between cell density and forest structure. High density cells with a basal area mean value of 45m2/ha were compared with low density cells with a basal area mean value of 18m2/ha. High density cells had a significantly higher species richness and higher diameter coefficient of variation than cells with low density. The density patterns in the canopy and understory layers were also found to be different, understory trees showing higher mortality rates with increasing tree size than canopy trees. Finally, either habitat heterogeneity or dispersal limitations are insufficient to explain maximum density patterns on their own. Our results show, however, that their joint effect may determine the formation of maximum density patterns in the two forests. Further studies in species-rich natural forests ecosystems are needed to explain density patterns in different ecosystems.

Chemical control of isolated invasive conifers using a novel aerial spot application method

SummaryConifer species, which have formed the foundation of commercial forestry industry in many countries, are known to be invasive in natural ecosystems, especially in the Southern Hemisphere. Controlling isolated invasive conifers before they reach reproductive maturity is an essential element of any strategy that aims to reduce spread rate of these species. Using a novel helicopter‐mounted spot‐application gun, which delivers a precise dosage to the crown of each tree, the objective of this research was to test the efficacy of three triclopyr‐based treatments against the four most vigorous wilding conifer species (Pinus contorta, Pinus nigra, Pinus sylvestris and Pseudotsuga menziesii) under New Zealand field conditions. Herbicides tested were triclopyr at two different rates and a combination of triclopyr and picloram. Treated trees covered a wide range of heights (c. 0.5–16 m), and measurements of mortality taken two years post‐herbicide application were used to examine variation in efficacy of the herbicides. Successful treatment was defined by a mortality rate of 85% or higher. A logistic regression model was fitted to the mortality data and used to derive threshold tree heights at which 85% mortality occurred, H85. For all four species, the most effective treatment was application of 1000 ml of herbicide mixture per tree that contained 120 and 20 g, respectively, of the active ingredients triclopyr and picloram. There was a significant decline in efficacy of this treatment with increases in tree size for all four species. Values of H85 for this treatment were 7.4 m for P. nigra, 8.3 m for P. menziesii, 9.7 m for P. contorta and >10 m for P. sylvestris. The methods developed here could be used to effectively manage emerging conifer infestations before they become problematic.

Allometric Biomass, Biomass Expansion Factor and Wood Density Models for the OP42 Hybrid Poplar in Southern Scandinavia

Biomass and biomass expansion factor functions are important in wood resource assessment, especially with regards to bioenergy feedstocks and carbon pools. We sampled 48 poplar trees in seven stands with the purpose of estimating allometric models for predicting biomass of individual tree components, stem-to-aboveground biomass expansion factors (BEF) and stem basic densities of the OP42 hybrid poplar clone in southern Scandinavia. Stand age ranged from 3 to 31 years, individual tree diameter at breast height (dbh) from 1.2 to 41 cm and aboveground tree biomass from 0.39 to 670 kg. Models for predicting total aboveground leafless, stem and branch biomass included dbh and tree height as predictor variables and explained more than 97 % of the total variation. The BEF was approaching 2.0 for the smallest trees but declined with increasing tree size and stabilized around 1.2 for trees with dbh >10 cm. Average stem basic density was more than 400 kg m−3 for the smallest trees but declined with increasing tree height and stabilized around 355 kg m−3, at a tree height of about 20 m. Existing biomass functions from the literature all underestimated the measured sample tree biomass. Possible explanations include not only differences in competition among trees in the examined stands and site conditions but also differences in sampling procedures. We observed that basic density increased with height above the ground. This trend may have led to the observed underestimation by existing biomass functions including only few samples from the lower end of the stems.

Modelling forage potential for red deer: a case study in post-disturbance young stands of rowan

Recently, the red deer ( Cervus elaphus ) population has increased considerably and caused serious damage in forest stands in Slovakia as well as in other Central-European countries. Rowan ( Sorbus aucuparia L.) is the tree species that is most intensively browsed and stripped by deer, especially during young stages of tree development. Our research focuses on estimating rowan mass consumption by red deer in young stands which developed after large-scale wind disturbance that occurred in the Tatra National Park in 2004. New models were developed for estimating the mass of tree components that are potentially edible by red deer using tree-base diameter as an independent variable. The results showed that the mass contribution of particular tree components to accessible deer forage depended on tree size (tree-base diameter). At stand level, total forage potential increased with an increase in tree size. However, whereas the quantity of bark available for stripping increased with tree size, the total mass accessible for browsing (leader shoot and branches with foliage) decreased. For instance, the contribution of stem bark to total forage potential in stands with a mean tree-base diameter of 20 mm and 50 mm was 15% and 50%, respectively. Theoretically, if all tree mass potential is consumed by red deer, young rowan stands (considering tree coverage of 50%) growing within an area of 100 m2 might provide sufficient forage for one adult deer for ca. 10 days. It is suggested that rowan species should not be removed from forest stands in territories with a high deer population in order to decrease the potential damage of other, commercially important, tree species.

Trees increase their P:N ratio with size.

Phosphorus (P) tends to become limiting in aging terrestrial ecosystems, and its resorption efficiency is higher than for other elements such as nitrogen (N). We thus hypothesized that trees should store more P than those other elements such as N when tree size increases and that this process should be enhanced in slow-growing late successional trees. Catalan forests. We have used data from the Catalan Forest Inventory that contains field data of the P and N contents of total aboveground, foliar and woody biomasses of the diverse Mediterranean, temperate and alpine forests of Catalonia (1018 sites). We used correlation and general lineal models (GLM) to analyze the allometric relationships between nutrient contents of different aboveground biomass fractions (foliar, branches and stems) and total aboveground biomass. Aboveground forest P content increases proportionally more than aboveground forest N content with increasing aboveground biomass. Two mechanisms underlie this. First, woody biomass increases proportionally more than foliar biomass having woody biomass higher P:N ratio than foliar biomass. Second, wood P:N ratio increases with tree size. These results are consistent with the generally higher foliar resorption of P than of N. Slow-growing species accumulate more P in total aboveground with size than fast-growing species mainly as a result of their large capacity to store P in wood. Trees may have thus developed long-term adaptive mechanisms to store P in biomass, mainly in wood, thereby slowing the loss of P from the ecosystems, reducing its availability for competitors, and implying an increase in the P:N ratio in forest biomass with aging. This trend to accumulate more P than N with size is more accentuated in slow-growing, large, long-living species of late successional stages. This way they partly counterbalance the gradual decrease of P in the soil.

Growth adjustments of conifers to drought and to century-long irrigation

Our knowledge on tree responses to drought is mainly based on short-term manipulation experiments which do not capture any possible long-term adjustments in this response. Therefore, historical water channels in inner-Alpine dry valleys were used as century-long irrigation experiments to investigate adjustments in tree growth to contrasting water supply. This involved quantifying the tree-ring growth of irrigated and non-irrigated (control) Scots pine (Pinus sylvestris L.) in Valais (Switzerland), as well as European larch (Larix decidua Mill.) and black pine (Pinus nigra Arnold) in Vinschgau (Italy). Furthermore, the adjustments in radial growth of Scots pine and European larch to an abrupt stop in irrigation were analyzed.Irrigation promoted the radial growth of all tree species investigated compared to the control: (1) directly through increased soil water availability, and (2) indirectly through increased soil nutrients and humus contents in the irrigated plots. Irrigation led to a full elimination of growth responses to climate for European larch and black pine, but not for Scots pine, which might become more sensitive to drought with increasing tree size in Valais. For the control trees, the response of the latewood increment to water availability in July/August has decreased in recent decades for all species, but increased in May for Scots pine only. The sudden irrigation stop caused a drop in radial growth to a lower level for Scots pine or similar level for larch compared to the control for up to ten years. However, both tree species were then able to adjust to the new conditions and subsequently grew with similar (Scots pine) or even higher growth rates (larch) than the control.To estimate the impact of climate change on future forest development, the duration of manipulation experiments should be on longer time scales in order to capture adjustment processes and feedback mechanisms of forest ecosystems.

Neighborhood analysis of underplanted Korean pine demography in larch plantations: Implications for uneven-aged management in northeast China

Quantifying the growth and survival of underplanted Korean pine (Pinus koraiensis Sieb. et Zucc.) in existing plantations provides a critical step toward developing an uneven-aged management system in northeast China. We conducted neighborhood analysis to quantify the effect of individual-level competition on the growth and survival of underplanted Korean pines in larch plantations (Larix olgensis A. Henry and Larix kaempferi [Lamb.] Carr.). The growth rate depended significantly on target tree size and the strength of competition from neighboring trees, whereas survival depended solely on tree size. The survival rate increased sharply with increasing tree size, and trees that had reached ca. 3cm in diameter at a height of 0.3m rarely died, even under strong competition. The effect of competition from neighbors on growth was determined by the neighboring tree’s size and its spatial relationship with the target tree. Neighbors located near and southwest of the target tree produced stronger competition than trees at greater distances and different azimuths. Our simulation in which Korean pines were underplanted in strip openings showed that their growth rates increased with increasing strip width. The growth rate was also related to the strip’s azimuth and the underplanting position within the opening. The results indicate that planting Korean pines in northwest to southeast strips would create a more heterogeneous size structure than planting in northeast to southwest strips. Our results suggest that spatial factors such as strip width, strip direction, and planting position strongly influence the development and productivity of uneven-aged Korean pine–larch stands.

Is there a ‘browse trap’? Dynamics of herbivore impacts on trees and grasses in an African savanna

Summary Despite widespread acknowledgement that large mammal herbivory can strongly affect vegetation structure in savanna, we still lack a theoretical and practical understanding of savanna dynamics in response to herbivory. Like fire, browsing may impose height‐structured recruitment limitations on trees (i.e. a ‘browse trap’), but the demographics of herbivore effects have rarely been considered explicitly. Evidence that cohorts of trees in savannas may establish during herbivore population crashes and persist long term in savanna landscapes is anecdotal. Here, we use an experimental approach in Hluhluwe iMfolozi Park in South Africa, examining the response of grass biomass and tree populations to 10 years of graduated herbivore exclusion, and their subsequent response when exclosures were removed. We found that grazer exclusion increased grass biomass and that, despite presumable increases in fire intensity and grass competition, herbivore – especially mesoherbivore, including impala and nyala – exclusion resulted in increases in tree size. After herbivore reintroduction, grazers reduced grass biomass over short time‐scales, but tree release from browsing persisted, regardless of tree size. Synthesis. This work provides the first experimental evidence that release from browsing trumps grazer–grass–fire interactions to result in increases in tree size that persist even after browser reintroduction. Escape from the ‘browse trap’ may be incremental and not strictly episodic, but, over longer time‐scales, reductions in browsing pressure may lead to tree establishment events in savanna that persist even during periods of intense browsing. Explicitly considering the temporal demographic effects of browsing will be the key for a much‐needed evaluation of the potential global extent of herbivore impacts in savanna.

Linking functional traits and demographic rates in a subtropical tree community: the importance of size dependency

Summary An important goal in plant community ecology is to understand how species traits determine demographic performance. Several functional traits have been shown to correlate with growth and mortality rates in trees, but less is known about how the relationships between functional traits and demographic rates change with tree size. We examined the associations of functional traits with growth and mortality across 43 tree species in the Fushan 25‐ha subtropical rain forest plot in northern Taiwan. We estimated the 95th percentile maximum stem diameter, wood density and six leaf functional traits (leaf area, specific leaf area, thickness, succulence, and mass‐based nitrogen and phosphorus contents) obtained from leaves on juvenile and adult individuals of each species. To quantify size‐dependent changes in growth and mortality, relative growth rate (RGR) and mortality were estimated as a function of stem diameter using hierarchical Bayesian models. These rate estimates were then correlated with functional traits at a range of stem diameter classes. Relationships between functional traits and demographic rates varied with tree size. Maximum size was positively correlated with RGR across a wide range of tree sizes. Wood density was negatively correlated with RGR and mortality for small‐sized trees. Leaf traits such as leaf area and specific leaf area at juvenile and adult stages were associated more strongly with demographic rates for corresponding sizes than from other sizes. Synthesis. The observed size‐dependent changes in the trait–demography relationships are possibly due to the effects of developmental and environmental changes with increasing tree size. The underlying effects of functional traits on demographic performance vary with tree size, and this should influence dynamics in a tree community.

Smooth bark surfaces can defend trees against insect attack: resurrecting a ‘slippery’ hypothesis

SummarySmooth bark on trees and shrubs was historically hypothesized to be an anatomical defence against epiphytic vegetation and phytophagous insects. This hypothesis has fallen from favour, yet no clear tests of bark texture as a defence against insects have been published.We tested the smooth bark defence hypothesis using bark beetles specialized in attacking pine trees as model insects, andPinus flexilis(limber pine) – a widespread tree that can have both smooth and rough bark surfaces on the same stem – as the model tree. We investigated the effects of bark texture on the locations of bark beetle attacks on trees with a combination of field surveys and experiments in the Colorado Rocky Mountains, USA.Bark beetle attacks were overwhelmingly located on rough bark surfaces and virtually absent from smooth bark. Increasing proportional coverage by smooth bark was negatively related to bark beetle attacks per square metre of bark surface. Experimental tests of bark beetles' ability to grip smooth versus rough bark revealed that bark beetles have difficulty gripping and quickly fell from smooth bark but not from rough bark.Smooth bark was negatively related to increasing tree size, but our models indicated that even partial coverage by smooth bark on a tree's trunk can significantly reduce total bark beetle attacks – this reduction likely improves tree fitness as bark beetles must aggregate to overcome tree defences.Synthesis. Our results indicate that smooth bark on trees can act as an anatomical defence against insects by reducing their ability to grip a tree's surface – even for insects specialized in attacking tree stems. Similar to other forms of anti‐insect defence (i.e. secondary chemistry, leaf toughness), smooth bark appears to be influenced by plant ontogeny whereby younger trees have greater defences than older trees. Understanding the adaptive significance of bark texture will require continued field and genetic study. Nevertheless, our results revealed that smooth bark texture increases tree resistance to phytophagous insects calling for the resurrection and vetting of the smooth bark defence hypothesis.

Linking size‐dependent growth and mortality with architectural traits across 145 co‐occurring tropical tree species

Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.

GPU 기반의 묶음 LOD 기법을 이용한 지형 렌더링의 가속화 기법

대용량 지형 데이터는 최신 그래픽 하드웨어를 사용해도 실시간으로 표현하기가 어렵다. 일반적으로 이런 대용량 지형 데이터를 실시간에 처리하기 위해 연속 상세 단계 기법 같은 메쉬 간략화 기법이 사용된다. 하지만 기하 분할(geometry splitting)과 같이 기존의 GPU기반 사진트리를 사용하는 기법은 트리의 깊이가 깊어질수록 많은 정점을 사용하고 이를 다시 재전송하기 때문에 성능이 저하되며 텍스처를 이용해 트리를 구성하기 때문에 트리의 용량이 커지는 단점이 있다. 이런 단점을 해결하기 위해 본 논문에서는 사진트리 기반의 묶음(chunk)으로 구성된 상세 단계 선별 기법을 GPU에서 처리하는 실시간 지형 렌더링 기법을 제안한다. 제안하는 방법은 기하 분할 기법의 단점을 해결하기 위해 트리 탐색을 제한하고 테셀레이터(tessellator)에서 묶음을 생성한 후 렌더링함으로써 트리 탐색을 줄이고 묶음을 GPU에서 바로 생성할 수 있어 효율적으로 지형을 렌더링할 수 있다. It is hard to represent massive terrain data in real-time even using recent graphics hardware. In order to process massive terrain data, mesh simplification method such as continuous Level-of-Detail is commonly used. However, existing GPU-based methods using quad-tree structure such as geometry splitting, produce lots of vertices to traverse the quad-tree and retransmit those vertices back to the GPU in each tree traversal. Also they have disadvantage of increase of tree size since they construct the tree structure using texture. To solve the problem, we proposed GPU-base chunked LOD technique for real-time terrain rendering. We restrict depth of tree search and generate chunks with tessellator in GPU. By using our method, we can efficiently render the terrain by generating the chunks on GPU and reduce the computing time for tree traversal.

Effects of precommercial thinning on the forest value chain in northwestern New Brunswick: Part 3 – Incidence of root and butt decay

The Green River precommercial thinning (PCT) trials were established between 1959 and 1961 in naturally regenerating balsam fir (Abies balsamea [L.] Mill.)-dominated stands an average of eight years after overstory removal. Three nominal spacings of 4 ft (1.2 m), 6 ft (1.8 m) and 8 ft (2.4 m) were compared to an unthinned control in six replicate blocks. In the fall of 2008, following completion of the ninth sequential evaluation of the study’s 48 permanent sample plots, three of the six replicates were clearcut harvested; butt rot data were collected immediately afterwards. To date, forest management research and goals have focused on the benefits of PCT, such as increased tree size and merchantable volume, shorter rotation ages, and better stem form and uniformity. Comparatively little attention has been placed on negative aspects of PCT, such as the incidence and development of root and butt rots, and their impact on fibre yields and wood product values. Results from the Green River study provide evidence that PCT may increase the incidence of butt rot in balsam fir, with incidence proportional to thinning intensity (p < 0.01). We also observed incidence and volume of butt rot to increase with stem diameter (p ≤ 0.05). The experiment suggests that factors such as stand age at time of thinning, and age at the time of harvest are important considerations when it comes to mitigating the impacts of butt rot through forest management.

Age-dependent climate–growth relationships and regeneration ofPicea abiesin a drought-prone mixed-coniferous forest in the Alps

Within dry inner Alpine environments climate warming is expected to affect the development of forest ecosystems by changing species composition and inducing shifts in forest distribution. By applying dendroecological techniques we evaluated climate sensitivity of radial growth and establishment of Picea abies in a drought-prone mixed-coniferous forest in the Austrian Alps. Time series of annual increments were developed from > 220 trees and assigned to four age classes. While radial growth of old P. abies trees (mean age 121 and 174 yr) responded highly significant to May-June precipitation, young trees (mean age 28 and 53 yr) were insensitive to precipitation in the current year. Because tree age was closely correlated to height and diameter (r2 = 0.709 and 0.784, respectively), we relate our findings to the increase in tree size rather than age per se. Synchronicity found among trend in basal area increment and tree establishment suggests that canopy openings increased light and water availability, which favoured growth and establishment of moderately shade-tolerant P. abies. We conclude that although P. abies is able to regenerate at this drought prone site, increasing inter-tree competition for water in dense stands gradually lowers competitive strength and restricts scattered occurrence to dry-mesic sites.

Temperature‐induced responses of xylem structure of Larix sibirica (Pinaceae) from the Russian Altay

Xylem structure determines the hydraulic and mechanical properties of a stem, and its plasticity is fundamental for maintaining tree performance under changing conditions. Unveiling the mechanism and the range of xylem adjustment is thus necessary to anticipate climate change impacts on vegetation. To understand the mechanistic process and the functional impact of xylem responses to warming in a cold-limited environment, we investigated the relationship between temperature and tracheid anatomy along a 312-yr tree-ring chronology of Larix sibirica trees from the Altay Mountains in Russia. Climate-growth analyses indicated that warming favors wider earlywood cell lumen, thicker latewood walls, denser maximum latewood, and wider rings. The temperature signal of the latewood was stronger (r > 0.7) and covered a longer and more stable period (from June to August) than that of earlywood and tree-ring width. Long-term analyses indicated a diverging trend between lumen and cell wall of early- and latewood. Xylem anatomy appears to respond to warming temperatures. A warmer early-growing season raises water conduction capacity by increasing the number and size of earlywood tracheids. The higher-performing earlywood tracheids promote more carbon fixation of the latewood cells by incrementing the rate of assimilation when summer conditions are favorable for growth. The diverging long-term variation of lumen and cell wall in earlywood vs. latewood suggests that xylem adjustments in latewood increase mechanical integrity and support increasing tree size under the ameliorated growing conditions.

Production Trends in Mature Macadamia Orchards and the Effects of Selective Limb Removal, Side-hedging, and Topping on Yield, Nut Characteristics, Tree Size, and Economics

Yields of macadamia (Macadamia integrifolia, M. tetraphylla, and hybrids) orchards tend to increase with increasing tree size up to ≈94% light interception. Beyond this, there is some indication that crowding leads to yield decline, but the evidence is limited to one site. Increasing tree size and orchard crowding also present numerous management problems, including soil erosion, harvest delays, and increased pest and disease pressure. The aim of this study was to better characterize long-term yield trends in mature orchards and to assess the effects of manual and mechanical pruning strategies on yield, nut characteristics, tree size, and economics. We monitored yield at four sites in mature ‘344’ and ‘246’ orchards for up to seven years and confirmed a decline in yield with crowding for three of the sites. There was a small increase in yield over time at the fourth site, which may reflect the lower initial level of crowding and shorter monitoring period compared with the other sites, and highlights the need for long-term records to establish yield trends. Pruning to remove several large limbs from ‘246’ trees to improve light penetration into the canopy increased yield relative to control trees but the effect was short-lived and not cost-effective. Removal of a codominant leader from ‘344’ trees reduced yield by 21%. Annual side-hedging of ‘246’ trees reduced yield by 12% and mechanical topping of ‘344’ trees caused a substantial reduction in yield of up to 50%. Removal of limbs in the upper canopy to reduce the height of ‘344’ trees had less effect on yield than topping but re-pruning was not practical because of the extensive regrowth around the pruning cuts. Tree size control is necessary for efficient orchard management, but in this study, pruning strategies that controlled tree size also reduced yield. Research into the physiological response to pruning in macadamia is required to improve outcomes.

Scaling of xylem and phloem transport capacity and resource usage with tree size.

Xylem and phloem need to maintain steady transport rates of water and carbohydrates to match the exchange rates of these compounds at the leaves. A major proportion of the carbon and nitrogen assimilated by a tree is allocated to the construction and maintenance of the xylem and phloem long distance transport tissues. This proportion can be expected to increase with increasing tree size due to the growing transport distances between the assimilating tissues, i.e., leaves and fine roots, at the expense of their growth. We formulated whole tree level scaling relations to estimate how xylem and phloem volume, nitrogen content and hydraulic conductance scale with tree size, and how these properties are distributed along a tree height. Xylem and phloem thicknesses and nitrogen contents were measured within varying positions in four tree species from Southern Finland. Phloem volume, nitrogen amount and hydraulic conductance were found to be concentrated toward the branch and stem apices, in contrast to the xylem where these properties were more concentrated toward the tree base. All of the species under study demonstrated very similar trends. Total nitrogen amount allocated to xylem and phloem was predicted to be comparable to the nitrogen amount allocated to the leaves in small and medium size trees, and to increase significantly above the nitrogen content of the leaves in larger trees. Total volume, hydraulic conductance and nitrogen content of the xylem were predicted to increase faster than that of the phloem with increasing tree height in small trees (<~10 m in height). In larger trees, xylem sapwood turnover to heartwood, if present, would maintain phloem conductance at the same level with xylem conductance with further increases in tree height. Further simulations with a previously published xylem-phloem transport model demonstrated that the Münch pressure flow hypothesis could explain phloem transport with increasing tree height even for the tallest trees.

Partitioning of soil water among canopy trees during a soil desiccation period in a temperate mixed forest

Abstract. Complementary resource use is considered an important mechanism in the study of biodiversity effects. Here we explore how species identity, species mixture and tree size influence the vertical partitioning of soil water among canopy trees during a soil desiccation period. In the Hainich Forest, Germany, the species Fagus sylvatica, Tilia sp. and Fraxinus excelsior were studied in single- and three-species mixed clusters, each consisting of three co-dominant trees situated within a larger mixed forest stand. Vertical soil water uptake depth was assessed by analyzing the hydrogen stable isotope composition (deuterium, δD) of water from depth intervals throughout the soil profile and in tree xylem water. For single species clusters, a mixing model suggested that Fagus distinctively drew water from soil depths of 0.3–0.5 m, Tilia from 0.3–0.5 m and 0.5–0.7 m and Fraxinus mainly used water from 0.5–0.7 m. In mixed clusters, the uptake patterns of Fagus and Tilia were similar to those of the single-species clusters (mainly uptake form 0.3–0.5 m), but Fraxinus showed a different uptake pattern. Fraxinus in mixture had a somewhat homogenously distributed uptake over the soil depths 0.2–0.7 m. For single species clusters, there was no correlation between main soil water uptake depth and tree diameter, irrespective of variations in tree size. In contrast, for mixed clusters there was a significant decrease in the main uptake depth with increasing tree size (P&lt;0.001, R2adj = 0.73), irrespective of species mix. In consequence, soil water partitioning was strongest where species were mixed and tree size varied. We further analyzed whether single and mixed-species clusters differed in the level of water uptake, e.g. due to complementarity, but our soil water budgeting did not indicate any such differences. A possible explanation might be that the volume of water used is predominantly governed by properties at the stand level, such as aerodynamic roughness, rather than by processes acting at the meter scale between neighbouring trees. With respect to application, we assume that the upcoming close-to-nature forestry approach for the area, which fosters mixed stands of heterogonous diameters, may result in enhanced complementarity in soil water uptake among canopy trees.

Performance of jack pine (Pinus banksiana) rooted cuttings from proliferated dwarf shoots versus seedlings 8 years after planting

Growth and tree form characteristics of jack pine (Pinus banksiana Lamb.) rooted cuttings propagated from proliferated dwarf shoots (PDS) were compared with seedlings in two field trials 8 years after establishment. Results indicated that jack pine rooted cuttings from PDS can grow as well as seedlings and maintain acceptable tree form. Rooted cuttings of progeny from the 22 top-ranking open-pollinated families in a seedling seed orchard of jack pine were 4.2% taller and 10% larger in diameter at breast height than commercial seedlings tested on the same sites, which indicates that rooted cuttings have potential in realizing genetic gains in jack pine tree improvement programs. Rooted cuttings increased the proportion of trees with normal branching characteristics and reduced the percentage of trees with excessive heavy branches in the Sault St. Marie trial, which had larger tree sizes. However, longer term monitoring (20 to 25 years) is needed to determine stability of jack pine rooted cuttings planted on sandy soil where wind throw may become a problem as tree size increases.

“Lifespan” of non-excavated holes in a primeval temperate forest: A 30 year study

Numerous forest organisms rely on non-excavated tree-holes, which are often limited in managed European woods. Holes’ supply depends on formation rates and persistence times. I use 30years of data from a primeval forest (strictly protected reserve, Białowieża National Park, Poland) to determine how long non-excavated holes persist, whether their persistence varies across forest types, tree species and conditions, or bird species using them. From 1979 to 2010, I followed the fate of 1794 holes, used by 14 non-excavator bird species. Almost half of them were destroyed during this time, most often (40%) due to tree fall or break off of the hole-bearing section, growing over injury (28%) or decay of walls (24%). Holes were retained for a median of 12years, longer in deciduous habitats (11–13years) than in coniferous forests (4.4years). These differences were due to different sets of tree species used in different habitats. Lifespan of holes varied by tree species, ranging from 5 (Populus tremula) to 16.5years (Quercus robur). Persistence was much lower for holes in dead (5years) than in living (14years) substrates. It increased with increasing tree size. Differences in persistence times of holes used by various bird species was mostly a by-product of them selecting trees of different qualities. Holes in large living trees, with relatively hard wood, persisted longest. Their retention should become a conservation priority. Current forestry policies should be modified, to assure that hole-bearing trees are retained, and trees in which replacement holes could be formed are maintained.