Canopy structure, which can be defined as the sum of the sizes, shapes and relative placements of the tree crowns in a forest stand, is central to all aspects of forest ecology. But there is no accepted method for deriving canopy structure from the sizes, species and biomechanical properties of the individual trees in a stand. Any such method must capture the fact that trees are highly plastic in their growth, forming tessellating crown shapes that fill all or most of the canopy space.
We introduce a new, simple and rapidly-implemented model–the Ideal Tree Distribution, ITD–with tree form (height allometry and crown shape), growth plasticity, and space-filling, at its core. The ITD predicts the canopy status (in or out of canopy), crown depth, and total and exposed crown area of the trees in a stand, given their species, sizes and potential crown shapes. We use maximum likelihood methods, in conjunction with data from over 100,000 trees taken from forests across the coterminous US, to estimate ITD model parameters for 250 North American tree species. With only two free parameters per species–one aggregate parameter to describe crown shape, and one parameter to set the so-called depth bias–the model captures between-species patterns in average canopy status, crown radius, and crown depth, and within-species means of these metrics vs stem diameter. The model also predicts much of the variation in these metrics for a tree of a given species and size, resulting solely from deterministic responses to variation in stand structure.
This new model, with parameters for US tree species, opens up new possibilities for understanding and modeling forest dynamics at local and regional scales, and may provide a new way to interpret remote sensing data of forest canopies, including LIDAR and aerial photography.
The relationship between canopy structure and light transmission to the forest floor is of particular interest for studying the effects of succession, timber harvest, and silviculture prescriptions on understory plants and trees. Indirect measurements of leaf area index (LAI) estimated using gap fraction analysis with linear and hemispheric sensors have been commonly used to assess radiation interception by the canopy, although the two methods often yield inconsistent results. We compared simultaneously obtained measurements of LAI from a linear ceptometer and digital hemispheric photography in 21 forest stands on Prince of Wales Island, Alaska. We assessed the relationship between these estimates and allometric LAI based on tree diameter at breast height (LAIDBH). LAI values measured at 79 stations in thinned, un-thinned controls, old-growth and clearcut stands were highly correlated between the linear sensor (AccuPAR) and hemispheric photography, but the latter was more negatively biased compared to LAIDBH. In contrast, AccuPAR values were more similar to LAIDBH in all stands with basal area less than 30 m2ha−1. Values produced by integrating hemispheric photographs over the zenith angles 0–75° (Ring 5) were highly correlated with those integrated over the zenith angles 0–60° (Ring 4), although the discrepancies between the two measures were significant. On average, the AccuPAR estimates were 53% higher than those derived from Ring 5, with most of the differences in closed canopy stands (unthinned controls and old-growth) and less so in clearcuts. Following typical patterns of canopy closure, AccuPAR LAI values were higher in dense control stands than in old-growth, whereas the opposite was derived from Ring 5 analyses. Based on our results we advocate the preferential use of linear sensors where canopy openness is low, canopies are tall, and leaf distributions are clumped and angles are variable, as is common in the conifer forests of coastal Alaska.
The dependence of aboveground biomass and productivity of tropical forests on soil fertility is not fully understood, since previous studies yielded contrasting results. Here, we quantify aboveground biomass (AGB) and stem wood production, and examine the impact of soil chemistry on these parameters in mature tropical forest stands of the equatorial Andes in Ecuador. In 80 plots of 0.04 ha at four elevation levels (500, 1,000, 1,500 and 2,000 m a.s.l., total sample area = 3.2 ha), we measured ten important soil chemical parameters, inventoried all trees ≥10 cm dbh and monitored stem diameter growth with dendrometer tapes in 32 plots. Top canopy height and stem density significantly decreased from 500 to 2,000 m, while tree basal area increased and AGB remained invariant (344 ± 17 Mg DM ha−1, mean ± SE) with elevation. Wood specific gravity (WSG) showed a significant, but small, decrease. Stem wood production decreased from 4.5 to 3.2 Mg DM ha−1 year−1 along the transect, indicating a higher biomass turnover at lower elevations. The only soil variable that covaried with AGB was exchangeable K in the topsoil. WSG increased with decreases in N mineralisation rate, soil pH and extractable Ca and P concentrations. Structural equation modelling (SEM) revealed that nitrogen availability acts on stem wood production only indirectly through a negative relation between N mineralisation rate and WSG, and a positive effect of a lowered WSG on stem growth. The SEM analysis showed neither direct nor indirect effects of resin-extractable P on wood production, but a negative P influence on AGB. We conclude that nitrogen availability significantly influences productivity in these Andean forests, but both N and P are affecting wood production mainly indirectly through alterations in WSG and stem density; the growth-promoting effect of N is apparently larger than that of P.
Electronic supplementary material
The online version of this article (doi:10.1007/s00442-012-2295-y) contains supplementary material, which is available to authorized users.
Aboveground biomass; Ecuador; Soil nutrients; Tree growth; Wood production
Tree species differ from one another in, and display trade-offs among, a wide range of attributes, including canopy and understorey growth and mortality rates, fecundity, height and crown allometry, and crown transmissivity. But how does this variation affect the outcome of interspecific competition and hence community structure? We derive criteria for the outcome of competition among tree species competing for light, given their allometric and life-history parameters. These criteria are defined in terms of a new simple whole life-cycle measure of performance, which provides a simple way to organize and understand the many ways in which species differ. The general case, in which all parameters can differ between species, can produce coexistence, founder control or competitive exclusion: thus, competition for light need not be hierarchical as implied by R* theory. The special case in which species differ only in crown transmissivity produces neutral dynamics. The special case in which species differ in all parameters except crown transmissivity gives hierarchical competition, where the equivalent of R* is Zˆ*, the height at which trees enter the canopy in an equilibrium monoculture.
biomechanics; succession; community ecology; forest dynamics; pairwise invasibility; Sortie
Few studies have quantified regional variation in tree mortality, or explored whether species compositional changes or within-species variation are responsible for regional patterns, despite the fact that mortality has direct effects on the dynamics of woody biomass, species composition, stand structure, wood production and forest response to climate change. Using Bayesian analysis of over 430,000 tree records from a large eastern US forest database we characterised tree mortality as a function of climate, soils, species and size (stem diameter). We found (1) mortality is U-shaped vs. stem diameter for all 21 species examined; (2) mortality is hump-shaped vs. plot basal area for most species; (3) geographical variation in mortality is substantial, and correlated with several environmental factors; and (4) individual species vary substantially from the combined average in the nature and magnitude of their mortality responses to environmental variation. Regional variation in mortality is therefore the product of variation in species composition combined with highly varied mortality-environment correlations within species. The results imply that variation in mortality is a crucial part of variation in the forest carbon cycle, such that including this variation in models of the global carbon cycle could significantly narrow uncertainty in climate change predictions.
We examined the effect of competition on stem growth of Picea glauca and Populus tremuloides in boreal mixedwood stands during the stem exclusion stage. We combined traditional approaches of collecting competition data with dendrochronology to provide retrospective measurements of stem diameter growth. Several competition indices including stand basal area (BA), the sum of stem diameter at breast height (SDBH), and density (N) for the broadleaf and coniferous species, as well as similar indices considering only trees with diameters greater than each subject (BAGR, SDBHGR, and NGR), were evaluated. We used a nonlinear mixed model to characterize the basal area increment over the past 5, 10, 15, 20, 25, 30, and 35 years as a function of growth of nearby dominant trees, the size of the subject trees, deciduous and coniferous competition indices, and ecoregions. SDBHGR and BAGR were better predictors for spruce, and SDBHGR and NGR were better for aspen, respectively, than other indices. Results showed strongest correlations with long-term stem growth, as the best models integrated growth for 10–25 years for aspen and ≥25 for spruce. Our model demonstrated a remarkable capability (adjusted R2>0.67) to represent this complex variation in growth as a function of site, size and competition.
Background and Aims
Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.
Focus is given to relationships between tree height and top-shoot traits, which include the shoot's leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot's stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.
Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.
In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.
Allometry; current-year leader shoot; hierarchical Bayesian model; pipe model; tree height; water stress
Background and Aims
Our knowledge about the influences of environmental factors on tree growth is principally based on the study of dominant trees. However, tree social status may influence intra-annual dynamics of growth, leading to differential responses to environmental conditions. The aim was to determine whether within-stand differences in stem diameters of trees belonging to different crown classes resulted from variations in the length of the growing period or in the rate of cell production.
Cambial activity was monitored weekly in 2006 for three crown classes in a 40-year-old silver-fir (Abies alba) plantation near Nancy (France). Timings, duration and rate of tracheid production were assessed from anatomical observations of the developing xylem.
Cambial activity started earlier, stopped later and lasted longer in dominant trees than in intermediate and suppressed ones. The onset of cambial activity was estimated to have taken 3 weeks to spread to 90 % of the trees in the stand, while the cessation needed 6 weeks. Cambial activity was more intense in dominant trees than in intermediate and suppressed ones. It was estimated that about 75 % of tree-ring width variability was attributable to the rate of cell production and only 25 % to its duration. Moreover, growth duration was correlated to tree height, while growth rate was better correlated to crown area.
These results show that, in a closed conifer forest, stem diameter variations resulted principally from differences in the rate of xylem cell production rather than in its duration. Tree size interacts with environmental factors to control the timings, duration and rate of cambial activity through functional processes involving source–sink relationships principally, but also hormonal controls.
Cambial activity; forest-stand structure; silver fir (Abies alba); tree-ring formation; tree-to-tree competition; social status; wood anatomy; xylem cell differentiation
Background and Aims
Morphogenetic plasticity may be as important as physiological plasticity in determining plant adaptability to changing environmental conditions. This study examines the importance of crown plasticity of trees in stands.
A three-dimensional forest simulator is used to explore the impact of crown shape plasticity on tree growth. Crown deformation is mediated through the local response to light and overall allometric constraints governing tree dimensions. By altering shape response parameters of Hevea brasiliensis the impact of increased or decreased plasticity is explored in a variety of competitive environments defined by various combinations of tree density and relative frequency of different strategies. The possible interactions between plasticity and growth rate and plasticity and below-ground competition are also explored.
Crown plasticity confers competitive superiority in all cases studied. Interactions with other processes may downplay or enhance this competitive advantage.
Simulation results strongly suggest that crown plasticity does have a significant impact on tree performance in nature and that commonly observed crown shape deformation response of trees is of adaptive value.
Crown plasticity; 3D simulation; individual-based model; competition
A recent study explored the adaptive significance of trunk inclination for trees growing on steep slopes. The authors used an optimality argument to predict how much a tree should respond to sidelight. Their calculations of the costs of leaning are flawed, because: (i) leaf mass has an allometric relationship with total volume, rather than being a fixed proportion of wood mass; (ii) the cost of support wood tissue is mainly a growth rate cost, not a maintenance respiration cost; (iii) small trees are sufficiently elastic to need very little support tissue, and thus they have a different risk structure; and (iv) most crown gaps are ephemeral rather than permanent, which also changes the risk equation. The argument I proposed in a previous study, addressing exactly the same question, is that canopy species are under selective pressure to maintain a strong central trunk that will reach the canopy and thus should not respond to sidelight. A reproductive value model is presented to illustrate this evolutionary question. Small, short-lived species or those from habitats with permanent openings (such as river margins) should be responsive. Both my previous data and the data of the other study support my model, but the model in the other study leaves many facts unexplained.
Trunk Inclination Leaf Biomass Shade-Tolerance
Background and Aims
Changes in size inequality in tree populations are often attributed to changes in the mode of competition over time. The mode of competition may also fluctuate annually in response to variation in growing conditions. Factors causing growth rate to vary can also influence competition processes, and thus influence how size hierarchies develop.
Detailed data obtained by tree-ring reconstruction were used to study annual changes in size and size increment inequality in several even-aged, fire-origin jack pine (Pinus banksiana) stands in the boreal shield and boreal plains ecozones in Saskatchewan and Manitoba, Canada, by using the Gini and Lorenz asymmetry coefficients.
The inequality of size was related to variables reflecting long-term stand dynamics (e.g. stand density, mean tree size and average competition, as quantified using a distance-weighted absolute size index). The inequality of size increment was greater and more variable than the inequality of size. Inequality of size increment was significantly related to annual growth rate at the stand level, and was higher when growth rate was low. Inequality of size increment was usually due primarily to large numbers of trees with low growth rates, except during years with low growth rate when it was often due to small numbers of trees with high growth rates. The amount of competition to which individual trees were subject was not strongly related to the inequality of size increment.
Differences in growth rate among trees during years of poor growth may form the basis for development of size hierarchies on which asymmetric competition can act. A complete understanding of the dynamics of these forests requires further evaluation of the way in which factors that influence variation in annual growth rate also affect the mode of competition and the development of size hierarchies.
Annual growth rate; asymmetric competition; competition index; dendroecology; Gini coefficient; jack pine; Lorenz asymmetry coefficient; volume increment
The investigation was conducted in a savanna area covered by what was considered an undesirably dense stand of Colophospermum mopane trees, mainly because such a dense stand of trees often results in the suppression of herbaceous plants. The objectives of this study were to determine the influence of intensity of tree thinning on the dry matter yield of herbaceous plants (notably grasses) and to investigate differences in herbaceous species composition between defined subhabitats (under tree canopies, between tree canopies and where trees have been removed). Seven plots (65 × 180 m) were subjected to different intensities of tree thinning, ranging from a totally cleared plot (0 %) to plots thinned to the equivalent of 10 %, 20%, 35 %, 50% and 75 % of the leaf biomass of a control plot (100 %) with a tree density of 2711 plants ha-1. The establishment of herbaceous plants (grasses and forbs) in response to reduced competition from the woody plants was measured during three full growing seasons following the thinning treatments.
The grass component reacted positively to the tree thinning in terms of total dry matter (DM) yield, but forbs were negatively influenced. Rainfall interacted with tree density and the differences between grass DM yields in thinned plots during years of below average rainfall were substantially higher than those of the control. At high tree densities, yields differed little between seasons of varying rainfall. The relation between grass DM yield and tree biomass was curvilinear, best described by the exponential regression equation. Subhabitat differentiation by C. mopane trees did provide some qualitative benefits, with certain desirable grass species showing a preference for the subhabitat under tree canopies.
While it can be concluded from this study that high tree densities suppress herbaceous production, the decision to clear/thin the C. mopane trees should include additional considerations. Thinning of C. mopane with the exclusive objective of increasing productivity of the grass layer would thus invariably involve a compromise situation where some trees should be left for the sake of the qualitative benefits on the herbaceous layer, soil enrichment, provision of browse and stability of the ecosystem.
Background and Aims
The dynamical system of plant growth GREENLAB was originally developed for individual plants, without explicitly taking into account interplant competition for light. Inspired by the competition models developed in the context of forest science for mono-specific stands, we propose to adapt the method of crown projection onto the x–y plane to GREENLAB, in order to study the effects of density on resource acquisition and on architectural development.
The empirical production equation of GREENLAB is extrapolated to stands by computing the exposed photosynthetic foliage area of each plant. The computation is based on the combination of Poisson models of leaf distribution for all the neighbouring plants whose crown projection surfaces overlap. To study the effects of density on architectural development, we link the proposed competition model to the model of interaction between functional growth and structural development introduced by Mathieu (2006, PhD Thesis, Ecole Centrale de Paris, France).
Key Results and Conclusions
The model is applied to mono-specific field crops and forest stands. For high-density crops at full cover, the model is shown to be equivalent to the classical equation of field crop production (
Howell and Musick, 1985, in Les besoins en eau des cultures; Paris: INRA Editions). However, our method is more accurate at the early stages of growth (before cover) or in the case of intermediate densities. It may potentially account for local effects, such as uneven spacing, variation in the time of plant emergence or variation in seed biomass. The application of the model to trees illustrates the expression of plant plasticity in response to competition for light. Density strongly impacts on tree architectural development through interactions with the source–sink balances during growth. The effects of density on tree height and radial growth that are commonly observed in real stands appear as emerging properties of the model.
Functional–structural plant models; GREENLAB; competition for light; Beer–Lambert Law; plant plasticity; dynamical system
Tree-ring width, wood density, anatomical structure and 13C/12C ratios expressed as δ13C-values of whole wood of Picea abies were investigated for trees growing in closed canopy forest stands. Samples were collected from the alpine Renon site in North Italy, the lowland Hainich site in Central Germany and the boreal Flakaliden site in North Sweden. In addition, Pinus cembra was studied at the alpine site and Pinus sylvestris at the boreal site. The density profiles of tree rings were measured using the DENDRO-2003 densitometer, δ13C was measured using high-resolution laser-ablation-combustion-gas chromatography-infra-red mass spectrometry and anatomical characteristics of tree rings (tracheid diameter, cell-wall thickness, cell-wall area and cell-lumen area) were measured using an image analyzer. Based on long-term statistics, climatic variables, such as temperature, precipitation, solar radiation and vapor pressure deficit, explained <20% of the variation in tree-ring width and wood density over consecutive years, while 29–58% of the variation in tree-ring width were explained by autocorrelation between tree rings. An intensive study of tree rings between 1999 and 2003 revealed that tree ring width and δ13C-values of whole wood were significantly correlated with length of the growing season, net radiation and vapor pressure deficit. The δ13C-values were not correlated with precipitation or temperature. A highly significant correlation was also found between δ13C of the early wood of one year and the late wood of the previous year, indicating a carry-over effect of the growing conditions of the previous season on current wood production. This latter effect may explain the high autocorrelation of long-term tree-ring statistics. The pattern, however, was complex, showing stepwise decreases as well as stepwise increases in the δ13C between late wood and early wood. The results are interpreted in the context of the biochemistry of wood formation and its linkage to storage products. It is clear that the relations between δ13C and tree-ring width and climate are multi-factorial in seasonal climates.
Dendrochonology; Carbohydrate storage; Picea abies; Pinus cembra; Pinus sylvestris; Tracheid lumen area; Wood density
Tree mortality is a key process underlying forest dynamics and community assembly. Understanding how tree mortality is driven by simultaneous drivers is needed to evaluate potential effects of climate change on forest composition. Using repeat-measure information from c. 400,000 trees from the Spanish Forest Inventory, we quantified the relative importance of tree size, competition, climate and edaphic conditions on tree mortality of 11 species, and explored the combined effect of climate and competition. Tree mortality was affected by all of these multiple drivers, especially tree size and asymmetric competition, and strong interactions between climate and competition were found. All species showed L-shaped mortality patterns (i.e. showed decreasing mortality with tree size), but pines were more sensitive to asymmetric competition than broadleaved species. Among climatic variables, the negative effect of temperature on tree mortality was much larger than the effect of precipitation. Moreover, the effect of climate (mean annual temperature and annual precipitation) on tree mortality was aggravated at high competition levels for all species, but especially for broadleaved species. The significant interaction between climate and competition on tree mortality indicated that global change in Mediterranean regions, causing hotter and drier conditions and denser stands, could lead to profound effects on forest structure and composition. Therefore, to evaluate the potential effects of climatic change on tree mortality, forest structure must be considered, since two systems of similar composition but different structure could radically differ in their response to climatic conditions.
One of the challenge fruit growers are facing is to balance between tree production and vegetative growth from year to year. To investigate the existence of genetic determinism for reproductive behaviour in olive tree, we studied an olive segregating population derived from a cross between ‘Olivière’ and ‘Arbequina’ cultivars. Our strategy was based on (i) an annual assessment of individual trees yield, and (ii) a decomposition of adult growth units at the crown periphery into quantitative variables related to both flowering and fruiting process in relation to their growth and branching. Genetic models, including the year, genotype effects and their interactions, were built with variance function and correlation structure of residuals when necessary. Among the progeny, trees were either ‘ON’ or ‘OFF’ for a given year and patterns of regular vs. irregular bearing were revealed. Genotype effect was significant on yield but not for flowering traits at growth unit (GU) scale, whereas the interaction between genotype and year was significant for both traits. A strong genetic effect was found for all fruiting traits without interaction with the year. Based on the new constructed genetic map, QTLs with small effects were detected, revealing multigenic control of the studied traits. Many were associated to alleles from ‘Arbequina’. Genetic correlations were found between Yield and Fruit set at GU scale suggesting a common genetic control, even though QTL co-localisations were in spe`cific years only. Most QTL were associated to flowering traits in specific years, even though reproductive traits at GU scale did not capture the bearing status of the trees in a given year. Results were also interpreted with respect to ontogenetic changes of growth and branching, and an alternative sampling strategy was proposed for capturing tree fruiting behaviour. Regular bearing progenies were identified and could constitute innovative material for selection programs.
Standing dead trees are one component of forest ecosystem dead wood carbon (C) pools, whose national stock is estimated by the U.S. as required by the United Nations Framework Convention on Climate Change. Historically, standing dead tree C has been estimated as a function of live tree growing stock volume in the U.S.'s National Greenhouse Gas Inventory. Initiated in 1998, the USDA Forest Service's Forest Inventory and Analysis program (responsible for compiling the Nation's forest C estimates) began consistent nationwide sampling of standing dead trees, which may now supplant previous purely model-based approaches to standing dead biomass and C stock estimation. A substantial hurdle to estimating standing dead tree biomass and C attributes is that traditional estimation procedures are based on merchantability paradigms that may not reflect density reductions or structural loss due to decomposition common in standing dead trees. The goal of this study was to incorporate standing dead tree adjustments into the current estimation procedures and assess how biomass and C stocks change at multiple spatial scales.
Accounting for decay and structural loss in standing dead trees significantly decreased tree- and plot-level C stock estimates (and subsequent C stocks) by decay class and tree component. At a regional scale, incorporating adjustment factors decreased standing dead quaking aspen biomass estimates by almost 50 percent in the Lake States and Douglas-fir estimates by more than 36 percent in the Pacific Northwest.
Substantial overestimates of standing dead tree biomass and C stocks occur when one does not account for density reductions or structural loss. Forest inventory estimation procedures that are descended from merchantability standards may need to be revised toward a more holistic approach to determining standing dead tree biomass and C attributes (i.e., attributes of tree biomass outside of sawlog portions). Incorporating density reductions and structural loss adjustments reduces uncertainty associated with standing dead tree biomass and C while improving consistency with field methods and documentation.
carbon accounting; forest inventory; greenhouse gas; dead wood; snag; standing dead
Japanese black bears, a large-bodied omnivore, frequently create small gaps in the tree crown during fruit foraging. However, there are no previous reports of black bear-created canopy gaps. To characterize physical canopy disturbance by black bears, we examined a number of parameters, including the species of trees in which canopy gaps were created, gap size, the horizontal and vertical distribution of gaps, and the size of branches broken to create gaps. The size of black bear-created canopy gaps was estimated using data from branches that had been broken and dropped on the ground.
The disturbance regime was characterized by a highly biased distribution of small canopy gaps on ridges, a large total overall gap area, a wide range in gap height relative to canopy height, and diversity in gap size. Surprisingly, the annual rate of bear-created canopy gap formation reached 141.3 m2 ha–1 yr–1 on ridges, which were hot spots in terms of black bear activity. This rate was approximately 6.6 times that of tree-fall gap formation on ridges at this study site. Furthermore, this rate was approximately two to three times that of common tree-fall gap formation in Japanese forests, as reported in other studies.
Our findings suggest that the ecological interaction between black bears and fruit-bearing trees may create a unique light regime, distinct from that created by tree falls, which increases the availability of light resources to plants below the canopy.
Animal–plant interaction; Bear shelf; Canopy disturbance; Gap distribution; Gap formation; Gap height; Topography
Individual trees have been shown to exhibit strong relationships between DBH, height and volume. Often such studies are cited as justification for forest volume or standing biomass estimation through remote sensing. With resolution of common satellite remote sensing systems generally too low to resolve individuals, and a need for larger coverage, these systems rely on descriptive heights, which account for tree collections in forests. For remote sensing and allometric applications, this height is not entirely understood in terms of its location. Here, a forest growth model (SERA) analyzes forest canopy height relationships with forest wood volume. Maximum height, mean, H100, and Lorey's height are examined for variability under plant number density, resource and species. Our findings, shown to be allometrically consistent with empirical measurements for forested communities world-wide, are analyzed for implications to forest remote sensing techniques such as LiDAR and RADAR. Traditional forestry measures of maximum height, and to a lesser extent H100 and Lorey's, exhibit little consistent correlation with forest volume across modeled conditions. The implication is that using forest height to infer volume or biomass from remote sensing requires species and community behavioral information to infer accurate estimates using height alone. SERA predicts mean height to provide the most consistent relationship with volume of the height classifications studied and overall across forest variations. This prediction agrees with empirical data collected from conifer and angiosperm forests with plant densities ranging between 102–106 plants/hectare and heights 6–49 m. Height classifications investigated are potentially linked to radar scattering centers with implications for allometry. These findings may be used to advance forest biomass estimation accuracy through remote sensing. Furthermore, Lorey's height with its specific relationship to remote sensing physics is recommended as a more universal indicator of volume when using remote sensing than achieved using either maximum height or H100.
• Background and Aims Growth in trunk height in canopy openings is important for saplings. How saplings increase height growth in canopy openings may relate to crown architectural constraints. Responses of crown development to canopy openings in relation to trunk height growth were studied for saplings (0·2–2·5 m tall) of eight tropical submontane forest tree species in Indonesia. The results of this study were also compared with those of temperate trees in northern Japan.
• Methods The crown architecture differed among the eight tropical species, i.e. they had sparsely to highly developed branching structures. Crown allometry was compared among the eight species in each canopy condition (closed canopy or canopy openings), and between closed canopy and canopy openings within a species. A general linear regression model was used to analyse how each species increases height growth rate in canopy openings. Crown allometry and its plasticity were compared between tropical and temperate trees by a nested analysis of covariance.
• Key Results Tropical submontane trees had responses similar to cool-temperate trees, showing an increase in height in canopy openings, i.e. taller saplings of sparsely branched species increase height growth rates by increasing the sapling leaf area. Cool-temperate trees have a wider crown projection area and a smaller leaf area per crown projection area to avoid self-shading within a crown compared with tropical submontane trees. Plasticity of the crown projection area is greater in cool-temperate trees than in tropical submontane trees, probably because of the difference in leaf longevity.
• Conclusions This study concluded that interspecific variation in the responses of crown development to canopy openings in regard to increasing height related to the species' branching structure, and that different life-forms, such as evergreen and deciduous trees, had different crown allometry and plasticity.
Cool-temperate trees; crown allometry; crown architecture; height growth; Indonesia; saplings; plasticity; tropical trees
▶ We examined four individual tree models in Europe: BWIN, Moses, Silva and Prognaus. ▶ We simulated growth of open-grown trees and on research plots for 15 or 30 years. ▶ Height:diameter ratios were correctly predicted by all four models. ▶ Height:diameter ratios were within the bounds of open grown trees and dense stands. ▶ They decreased with age and density; dominant trees had lower ratios than mean trees.
Height:diameter ratios are an important measure of stand stability. Because of the importance of height:diameter ratios for forest management, individual-tree growth models should correctly depict height:diameter ratios. In particular, (i) height:diameter ratios should not exceed that of very dense stands, (ii) height:diameter ratios should not fall below that of open-grown trees, (iii) height:diameter ratios should decrease with increasing spacing, (iv) height:diameter ratios for suppressed trees should be higher than ratios for dominant trees. We evaluated the prediction of height:diameter ratios by running four commonly used individual-tree growth models in central Europe: BWIN, Moses, Silva and Prognaus. They represent different subtypes of individual-tree growth models, namely models with and without an explicit growth potential and models that are either distance-dependent (spatial) or distance-independent (non-spatial). Note that none of these simulators predict height:diameter ratios directly. We began by building a generic simulator that contained the relevant equations for diameter increment, height increment, and crown size for each of the four simulators. The relevant measures of competition, site characteristics, and stand statistics were also coded. The advantage of this simulator was that it ensured that no additional constraint was being imposed on the growth equations, and that initial conditions were identical. We then simulated growth for a 15- and 30-year period for Austrian permanent research plots in Arnoldstein and in Litschau, which represent stands at different age-classes and densities. We also simulated growth of open-grown trees and compared the results to the literature. We found that the general pattern of height:diameter ratios was correctly predicted by all four individual-tree growth models, with height:diameter ratios above that of open-grown trees and below that of very dense stands. All models showed a decrease of height:diameter ratios with age and an increase with stand density. Also, the height:diameter ratios of dominant trees were always lower than that of mean trees. Although in some cases the observed and predicted height:diameter ratios matched well, there were cases where discrepancies between observed and predicted height:diameter ratios would be unacceptable for practical management predictions.
Stand stability; Height:diameter ratio; Individual-tree growth model; Model evaluation; Open-grown trees
Tree growth models are supposed to contain stand growth laws as so called “emergent properties” which derive from interactions of individual-tree growth and mortality functions. This study investigates whether the evolving tree species composition in a long term simulation by the distance-independent tree growth model PrognAus matches the species composition of the potential natural vegetation type which is expected to occur if one refrains from further management interventions and major disturbances, climate change, and changes in site conditions can be excluded. For this purpose the development of 6933 sample plots of the Austrian National Forest Inventory was predicted for 2500 years. The resulting species proportions, derived from volume per hectare of 15 tree species or species groups, were used to classify every sample plot according to potential natural forest types, following a classification scheme based on expert knowledge. These simulated potential natural vegetation types were compared with expert reconstructions of the sample plots of the Austrian National Forest Inventory. A total of 5789 plots were actually classified with the scheme; in 33% of the cases the classification on the basis of the PrognAus-simulations was identical with the classification by the Austrian National Forest Inventory. A predominantly correct classification was achieved for the subalpine Picea abies-type and the Fagus sylvatica-type although PrognAus showed a tendency to overestimate the proportion of F. sylvatica and P. abies. Weaknesses in the ability to simulate forest types dominated by Quercus spp., Acer spp., and Pinus sylvestris were identified. This shortcoming might be caused by the mortality model which allows a larger diameter at breast height for F. sylvatica or by the ingrowth model whose terms for the consideration of inter-specific competition may lead to a disadvantage of Quercus spp., P. sylvestris, and Abies alba. Moreover, the ingrowth model might be influenced by management effects and the effect of selective browsing.
Individual-tree growth model; Species distribution; Potential natural vegetation type; Forest stand development
Background and Aims
There is good evidence for deciduous trees that competition for carbohydrates from shoot growth accentuates early fruit abscission and reduces yield but the effect for evergreen trees is not well defined. Here, whole-tree tip-pruning at anthesis is used to examine the effect of post-pruning shoot development on fruit abscission in the evergreen subtropical tree macadamia (Macadamia integrifolia, M. integrifolia × tetraphylla). Partial-tree tip-pruning is also used to test the localization of the effect.
In the first experiment (2005/2006), all branches on trees were tip-pruned at anthesis, some trees were allowed to re-shoot (R treatment) and shoots were removed from others (NR treatment). Fruit set and stem total non-structural carbohydrates (TNSC) over time, and yield were measured. In the second experiment (2006/2007), upper branches of trees were tip-pruned at anthesis, some trees were allowed to re-shoot (R) and shoots were removed from others (NR). Fruit set and yield were measured separately for upper (pruned) and lower (unpruned) branches.
In the first experiment, R trees set far fewer fruit and had lower yield than NR trees. TNSC fell and rose in all treatments but the decline in R trees occurred earlier than in NR trees and coincided with early shoot growth and the increase in fruit abscission relative to the other treatments. In the second experiment, fruit abscission on upper branches of R trees increased relative to the other treatments but there was little difference in fruit abscission between treatments on lower branches.
This study is the first to demonstrate an increase in fruit abscission in an evergreen tree in response to pruning. The effect appeared to be related to competition for carbohydrates between post-pruning shoot growth and fruit development and was local, with shoot growth on pruned branches having no effect on fruit abscission on unpruned branches.
Pruning; hedging; shoot growth; fruit set; abscission; canopy management; Macadamia integrifolia
Losses of plant species diversity can affect ecosystem functioning, with decreased primary productivity being the most frequently reported effect in experimental plant assemblages, including tree plantations. Less is known about the role of biodiversity in natural ecosystems, including forests, despite their importance for global biogeochemical cycling and climate. In general, experimental manipulations of tree diversity will take decades to yield final results. To date, biodiversity effects in natural forests therefore have only been reported from sample surveys or meta-analyses with plots not initially selected for diversity. We studied biomass and growth of subtropical forests stands in southeastern China. Taking advantage of variation in species recruitment during secondary succession, we adopted a comparative study design selecting forest plots to span a gradient in species richness. We repeatedly censored the stem diameter of two tree size cohorts, comprising 93 species belonging to 57 genera and 33 families. Tree size and growth were analyzed in dependence of species richness, the functional diversity of growth-related traits, and phylogenetic diversity, using both general linear and structural equation modeling. Successional age covaried with diversity, but differently so in the two size cohorts. Plot-level stem basal area and growth were positively related with species richness, while growth was negatively related to successional age. The productivity increase in species-rich, functionally and phylogenetically diverse plots was driven by both larger mean sizes and larger numbers of trees. The biodiversity effects we report exceed those from experimental studies, sample surveys and meta-analyses, suggesting that subtropical tree diversity is an important driver of forest productivity and re-growth after disturbance that supports the provision of ecological services by these ecosystems.
We investigate the spread of an exotic herb, Hieracium lepidulum, into a New Zealand Nothofagus forest with the aim of understanding how stand-development of tree populations, propagule pressure and invader persistence, affect invasion across the landscape and within communities. Using data repeatedly collected over 35 years, from 250 locations, we parametrize continuous-time Markov chain models and use these models to examine future projections of the invasion under a range of hypothetical scenarios. We found that the probability of invasion into a stand was relatively high following canopy disturbance and that local abundance of Hieracium was promoted by minor disturbances. However, model predictions extrapolated 45 years into the future show that neither the rate of landscape-level invasion, nor local population growth of Hieracium, was affected much by changing the frequency of canopy disturbance events. Instead, invasion levels were strongly affected by the ability of Hieracium to persist in the understorey following forest canopy closure, and by propagule supply from streams, forest edges and plants already established within the stand. Our results show that disturbance frequency has surprisingly little influence on the long-term trajectory of invasion, while invader persistence strongly determines invasion patterns.
Hieracium; invader persistence; Markov chain models; Nothofagus; propagule pressure; stand-development