Summer drought: a driver for crown condition and mortality of Norway spruce in Norway

Summer drought, i.e. unusually dry and warm weather, has been a significant stress factor for Norway spruce in southeast Norway during the 14 years of forest monitoring. Dry and warm summers were followed by increases in defoliation, discolouration of foliage, cone formation and mortality. The causal mechanisms are discussed. Most likely, the defoliation resulted from increased needle‐fall in the autumn after dry summers. During the monitoring period 1988–2001, southeast Norway was repeatedly affected by summer drought, in particular, in the early 1990s. The dataset comprised 455 ‘Forest officers’ plots’ with annual data on crown condition and mortality. Linear mixed models were used for estimation and hypothesis testing, including a variance–covariance structure for the handling of random effects and temporal autocorrelation.     

Hydraulic and stomatal adjustment of Norway spruce trees to environmental stress

A study of how the water conducting systems of 30-50-year-old Norway spruce (Picea abies (L.) Karst.) trees growing at three sites adjust to shade and waterlogging indicated that water relations characteristics varied with the life histories of the trees. Xylem was more efficient at conducting water and stomata were more sensitive to atmospheric evaporative demand in trees subjected to favorable growth conditions (control trees) than in trees growing in shade or waterlogged conditions. At the same soil water availability, shade-grown trees suffered more severely from water deficit than control trees. Under conditions of high atmospheric vapor pressure deficit, foliage of shade-grown trees exhibited low water potentials, as a result of low hydraulic conductance of the vascular system and inefficient stomatal control. Because of the increased internal resistance to water flow, more negative leaf water potentials (Psi(x)) must be reached to provide an adequate water supply to the foliage. It is concluded that dynamic water stress is one of the main causes of the continuing growth retardation in suppressed Norway spruce trees after their release from the overstory. Trees growing in waterlogged soil (bog-grown trees) were characterized by weak stomatal control, resulting in large water losses from the foliage. Although bog-grown trees exhibited uneconomical water use, they possessed mechanisms (e.g., osmotic adjustment) that allowed leaves to tolerate low Psi(x) while stomata remained open. Under conditions of sufficient soil water availability and moderate atmospheric vapor pressure deficit, soil-to-leaf conductance was highest in bog-grown trees (1.45 +/- 0.06 mmol m(-2) s(-1) MPa(-1)), followed by control and shade-grown trees (1.04 +/- 0.04 and 0.77 +/- 0.05 mmol m(-2) s(-1) MPa(-1), respectively). The lowest soil-to-leaf conductance (0.45 +/- 0.04 mmol m(-2) s(-1) MPa(-1)) was recorded in control trees at high atmospheric evaporative demand, and was probably caused by tracheid cavitation.     

Midday depression of net photosynthesis in the tropical rainforest tree Eperua grandiflora: contributions of stomatal and internal conductances,respiration and Rubisco functioning

High midday temperatures can depress net photosynthesis. We investigated possible mechanisms underlying this phenomenon in leaves of Eperua grandiflora (Aubl.) Benth. saplings. This tropical tree establishes in small gaps in the rainforest canopy where direct sunlight can raise midday temperatures markedly. We simulated this microclimate in a growth chamber by varying air temperature between 28 and 38 degrees C at constant vapor pressure. A decrease in stomatal conductance in response to an increase in leaf-to-air vapor pressure difference (deltaW) caused by an increase in leaf temperature (Tleaf) was the principal reason for the decrease in net photosynthesis between 28 and 33 degrees C. Net photosynthesis decreased further between 33 and 38 degrees C. Direct effects on mesophyll functioning and indirect effects through deltaW were of similar magnitude in this temperature range. Mitochondrial respiration during photosynthesis was insensitive to Tleaf over the investigated temperature range; it thus did not contribute to midday depression of net photosynthesis. Internal conductance for CO2 diffusion in the leaf, estimated by combined gas exchange and chlorophyll fluorescence measurements, decreased slightly with increasing Tleaf. However, the decrease in photosynthetic rate with increasing Tleaf was larger and thus the difference in CO2 partial pressure between the substomatal cavity and chloroplast was smaller, leading to the conclusion that this factor was not causally involved in midday depression. Carboxylation capacity inferred from the CO2 response of photosynthesis increased between 28 and 33 degrees C, but remained unchanged between 33 and 38 degrees C. Increased oxygenation of ribulose-1,5-bisphosphate relative to its carboxylation and the concomitant increase in photorespiration with increasing Tleaf were thus not compensated by an increase in carboxylation capacity over the higher temperature range. This was the principal reason for the negative effect of high midday temperatures on mesophyll functioning.     

华北落叶松幼态程度与细胞超微结构之间关系的研究Ⅱ．树冠上部与树冠下部的一年生枝条

扦插是林木的重要繁殖方法。扦插生根率与枝条的幼态程度之间具有明显的相关性。扦插失败的主要原因之一是没有彻底了解无性繁殖材料组织与细胞的优化机理，而不能采取有效的措施和方法。核移植的试验表明，决定幼态的因素存在于细胞质之中。细胞器的形态数量变化以及超微结构水平可以鉴定细胞和组织的幼态程度。华北落叶松是中国北方重要造林树种，作者曾对胚、３龄和１２龄树木一年生枝条的细胞超微结构，进行过一些比较研究，表明无性繁殖材料的幼态程度具有明显的年龄效应。本文借助透射电子显微镜，对华北落叶松同一树冠不同部位一年生枝条的细胞超微结构进行比较研究。结果是，上部树冠与下部树冠相比，细胞中的质体具有明显的基粒片层结构和丰富的脂滴，前质体较少。线粒体数量多，具有更发达的嵴，一些线粒体处于增殖的初期阶段。液泡以次生溶酶体的形式存在。胞间连丝数量多，其中一些可观察到精细的管状结构。由此可见，树冠上部一年生枝条细胞超微结构较树冠下部复杂，即其幼态程度较低。在同一株树木中，幼态程度具有明显的部位效应。     

小尺度森林郁闭度测定方法及应用

郁闭度作为反映树冠闭锁程度、树木利用生活空间程度、森林结构及森林生长状况、生长环境等的重要因子之一.郁闭度的快速、准确测定一直以来都是学者们探讨的热点,但是,目前的郁闭度测定方法大多在工作量、精度与成本间无法兼顾.因此,提出一种基于地理信息系统(GIS)缓冲区分析功能进行郁闭度定量求算方法(简称“缓冲区法”),通过与树冠投影法实测的郁闭度特征进行方差分析,结果表明,缓冲区法与树冠投影法实测的郁闭度和树冠垂直投影图形状之间均无显著差异.     

Fine root condition relates to visible crown damage in Norway spruce in acidified soils

Allochthonous Norway spruce stands in the Kysucké Beskydy Mts. (north‐western Slovakia) have been exposed to substantial acid deposition in the recent past and grow in acidified soil conditions with mean pH of about 4.0 in the topsoil. We selected 90 spruce trees representing 30 triplets of different crown status: healthy, stressed and declining to assess the relationship between crown and fine root status. Sequential coring and in‐growth bags were applied to each triplet to investigate fine root mass and growth in the soil depths of 0–10 and 10–20 cm. Fine root quantity (biomass and necromass), turnover (production over standing stock), morphological features (specific root length, root tip density) and chemical properties (Ca:Al molar ratio) were compared among the abovementioned health status categories. Living fine root biomass decreased with increasing stress, while the ratio of dead to living biomass increased. Annual fine root production decreased and specific root length increased in stressed trees when compared to healthy or declining trees, a situation which may be related to the position of trees in the canopy (healthy and declining – dominant, stressed – co‐dominant). The Ca:Al ratio decreased with increasing crown damage, indicating a decreased ability to filter out aluminium. In conclusion, fine root status appears to be linked to visible crown damage and can be used as a tree health indicator.     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

Inherited narrow crown form, harvest index and stem biomass production in Norway spruce, Picea abies

The effect of crown form on stem biomass production was investigated in an 18-19-year-old Norway spruce stand (Picea abies (L.) Karst.). The harvest index was 0.271 in pendula trees, which have a heritable narrow crown form, and 0.235 in normal-crowned trees and the dry weights of stem biomass were 2.57 kg and 3.37 kg, respectively. However, the production of stem biomass per crown projected area was more than twice as much in pendula trees as in normal-crowned trees. Results indicate that the crown form of pendula spruce is less plastic than that of normal-crowned spruce suggesting that such trees might maintain a high stemwood yield per unit ground area with increasing stand density. Because pendula spruce has a higher harvest index and a less flexible crown than normal-crowned spruce, it might be useful for crop tree ideotype breeding.     

Effects of crown length on indole acetic acid (IAA) amounts in cambial region tissues in lower and upper trunks of sugi cultivars (Cryptomeria japonica) in September

Indole acetic acid (IAA) was believed to be an important regulator in xylem formation in conifers. However, few studies have been done on the endogenous amount of IAA in cambial region tissues in sugi (Cryptomeria japonica) trees. In this study, we report the IAA amounts in the lower and upper trunk of mature sugi cultivars (Kumotoshi, Yaichi and Obiaka) in September, and the effects of growth traits on the IAA amounts. The effect of height position (lower or upper trunk) on IAA amounts was found to be larger than that of genetic variation (Kumotoshi, Yaichi or Obiaka) by two-way ANOVA. There was no significant interaction effect (genetic variation × height position). The IAA amounts of all trees varied from 3 to 42 ng/cm² at the lower trunk and 9 to 47 ng/cm² at the upper trunk. Crown length and distance from crown base had significant positive and negative effects on IAA amounts in the trunk of sugi cultivars, respectively. Distance from crown base had a larger effect on IAA amounts than crown length. In Yaichi, taller trees had larger IAA amounts at both the lower and upper trunk.     

Growth and mortality in underplanted tree seedlings in response to variations in canopy closure of Norway spruce stands

Six broadleaved tree species and Picea abies (L.) Karst. wereplanted under spruce plantations of varying densities, in Sweden.Treatments included control (994 stems ha^–1), dense (538stems ha^–1), sparse (294 stems ha^–1) and gap (0stems ha^–1) overstory treatments. There was an increasein height and diameter growth from control to sparse overstorytreatment of all underplanted tree species except for ash (Fraxinusexcelsior L.) and Norway maple (Acer platanoides L.). Site conditionsmay have hampered the growth of these species, as well as wildcherry (Prunus avium L.). Both oak (Quercus robur L.) and sprucehad greater growth in the gap treatment, relative to the othertreatments. Insecticide application did not influence seedlinggrowth or survival. The performance of beech (Fagus sylvaticaL.), lime (Tilia cordata Mill.), spruce and oak was consistentwith shade tolerance ranking. Beech and lime had a very highsurvival rate, even under the densest canopy. The growth andmortality of ash, maple and wild cherry differed significantlyfrom what was expected. This experiment demonstrated significantvariation in interspecific growth and mortality between sevenplanted tree species in relation to canopy density. Correctsite and species selection is crucial when underplanting inshelterwood systems.     

Estimating single-tree branch biomass of Norway spruce with terrestrial laser scanning using voxel-based and crown dimension features

Abstract

Many remote sensing-based methods estimating forest biomass rely on allometric biomass models for field reference data. Terrestrial laser scanning (TLS) has emerged as a tool for detailed data collection in forestry applications, and the methods have been proposed to derive, e.g. tree position, diameter-at-breast-height, and stem volume from TLS data. In this study, TLS-derived features were related to destructively sampled branch biomass of Norway spruce at the single-tree level, and the results were compared to conventional allometric models with field measured diameter and height. TLS features were derived following two approaches: one voxel-based approach with a detailed analysis of the interaction between individual voxels and each laser beam. The features were derived using voxels of size 0.1, 0.2, and 0.4 m, and the effect of the voxel size was assessed. The voxel-derived features were compared to features derived from crown dimension measurements in the unified TLS point cloud data. TLS-derived variables were used in regression models, and prediction accuracies were assessed through a Monte Carlo cross-validation procedure. The model based on 0.4 m voxel data yielded the best prediction accuracy, with a root mean square error (RMSE) of 32%. The accuracy was found to decrease with an increase in voxel size, i.e. the model based on the 0.1 m voxel yielded the lowest accuracy. The model based on crown measurements had an RMSE of 34%. The accuracies of the predictions from the TLS-based models were found to be higher than from conventional allometric models, but the improvement was relatively small.     

Relationships between crown projected area and components of above-ground biomass in Norway spruce trees in even-aged stands: Empirical results and their interpretation

Sample tree material was reanalyzed in order to study the relationships between horizontal crown projected area and components of above-ground biomass in Norway spruce (Picea abies (L.) Karst.) trees growing in even-aged stands. The needle mass of dominant trees increased linearly with the increase in crown projected area, but in co-dominant and dominanted trees the increase in needle mass levelled off toward larger crown projected areas. The branch mass of dominant and co-dominant trees accumulated faster than linearly with increasing crown projected area, whereas in dominated trees an approximately linear relationship existed between these two variables. The increase in needle and branch mass per unit increase in crown projected area was highest in dominant trees and decreased to co-dominant and dominated trees, i.e. with tree position in the canopy. The stem mass accumulated obviously faster than linearly and similarly in all tree classes with the increase in crown projected area. The narrow crown shape indicated a high density of all components of above-ground biomass per unit of crown projected area.     

关于河南省黄河上中游天然林保护的建议

主要阐述了河南省黄河上中游天保区的现状、实施范围、目标及天然林保护应采取的措施。河南省天保区位于小浪底水库上游 ,保护好现有天然林 ,从根本上解决黄河的问题 ,保障小浪底等水利枢纽工程长期发挥效能有着重要的意义     

Effects of elevated CO(2) concentration and nutrition on net photosynthesis,stomatal conductance and needle respiration of field-grown Norway spruce trees

To study the effects of elevated CO(2) on gas exchange, nonstructural carbohydrate and nutrient concentrations in current-year foliage of 30-year-old Norway spruce (Picea abies (L.) Karst.) trees, branches were enclosed in ventilated, transparent plastic bags and flushed with ambient air (mean 370 &mgr;mol CO(2) mol(-1); control) or ambient air + 340 &mgr;mol CO(2) mol(-1) (elevated CO(2)) during two growing seasons. One branch bag was installed on each of 24 selected trees from control and fertilized plots. To reduce the effect of variation among trees, results from each treated branch were compared with those from a control branch on the same whorl of the same tree. Elevated CO(2) increased rates of light-saturated photosynthesis on average by 55% when measured at the treatment CO(2) concentration. The increase was larger in shoots with high needle nitrogen concentrations than in shoots with low needle nitrogen concentrations. However, shoots grown in elevated CO(2) showed a decrease in photosynthetic capacity compared with shoots grown in ambient CO(2). When measured at the internal CO(2) concentration of 200 &mgr;mol CO(2) mol(-1), photosynthetic rates of branches in the elevated CO(2) treatments were reduced by 8 to 32%. The elevated CO(2) treatment caused a 9 to 20% reduction in carboxylation efficiency and an 18% increase in respiration rates. In response to elevated CO(2), starch, fructose and glucose concentrations in the needles increased on average 33%, whereas concentrations of potassium, nitrogen, phosphorus, magnesium and boron decreased. Needle nitrogen concentrations explained 50-60% of the variation in photosynthesis and CO(2) acclimation was greater at low nitrogen concentrations than at high nitrogen concentrations. We conclude that the enhanced photosynthetic rates found in shoots exposed to elevated CO(2) increased carbohydrate concentrations, which may have a negative feedback on the photosynthetic apparatus and stimulate cyanide-resistant respiration. We also infer that the decrease in nutrient concentrations of needles exposed to elevated CO(2) was the result of retranslocation of nutrients to other parts of the branch or tree.     

Sapwood temperature gradients between lower stems and the crown do not influence estimates of stand-level stem CO(2) efflux

Temperature plays a critical role in the regulation of respiration rates and is often used to scale measurements of respiration to the stand-level and calculate annual respiratory fluxes. Previous studies have indicated that failure to consider temperature gradients between sun-exposed stems and branches in the crown and shaded lower stems may result in errors when deriving stand-level estimates of stem CO(2) efflux. We measured vertical gradients in sapwood temperature in a mature lowland podocarp rain forest in New Zealand to: (1) estimate the effects of within-stem temperature variation on the vertical distribution of stem CO(2) efflux; and (2) use these findings to estimate stand-level stem CO(2) efflux for this forest. Large within-stem gradients in sapwood temperature (1.6 +/- 0.1 to 6.0 +/- 0.5 degrees C) were observed. However, these gradients did not significantly influence the stand-level estimate of stem CO(2) efflux in this forest (536 +/- 42 mol CO(2) ha(-1) day(-1)) or the vertical distribution of stem CO(2) efflux, because of the opposing effects of daytime warming and nighttime cooling on CO(2) efflux in the canopy, and the small fraction of the woody biomass in the crowns of forest trees. Our findings suggest that detailed measurements of within-stand temperature gradients are unlikely to greatly improve the accuracy of tree- or stand-level estimates of stem CO(2) efflux.     

Shoot growth and crown development: effect of crown position in three-dimensional simulations

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.     

Stand age and fine root biomass, distribution and morphology in a Norway spruce chronosequence in southeast Norway

We assessed the influence of stand age on fine root biomass and morphology of trees and understory vegetation in 10-, 30-, 60- and 120-year-old Norway spruce stands growing in sandy soil in southeast Norway. Fine root (< 1, 1-2 and 2-5 mm in diameter) biomass of trees and understory vegetation (< 2 mm in diameter) was sampled by soil coring to a depth of 60 cm. Fine root morphological characteristics, such as specific root length (SRL), root length density (RLD), root surface area (RSA), root tip number and branching frequency (per unit root length or mass), were determined based on digitized root data. Fine root biomass and morphological characteristics related to biomass (RLD and RSA) followed the same tendency with chronosequence and were significantly higher in the 30-year-old stand and lower in the 10-year-old stand than in the other stands. Among stands, mean fine root (< 2 mm) biomass ranged from 49 to 398 g m(-2), SLR from 13.4 to 19.8 m g(-1), RLD from 980 to 11,650 m m(-3) and RSA from 2.4 to 35.4 m(2) m(-3). Most fine root biomass of trees was concentrated in the upper 20 cm of the mineral soil and in the humus layer (0-5 cm) in all stands. Understory fine roots accounted for 67 and 25% of total fine root biomass in the 10- and 120-year-old stands, respectively. Stand age had no affect on root tip number or branching frequency, but both parameters changed with soil depth, with increasing number of root tips and decreasing branching frequency with increasing soil depth for root fractions < 2 mm in diameter. Specific (mass based) root tip number and branching density were highest for the finest roots (< 1 mm) in the humus layer. Season (spring or fall) had no effect on tree fine root biomass, but there was a small and significant increase in understory fine root biomass in fall relative to spring. All morphological characteristics showed strong seasonal variation, especially the finest root fraction, with consistently and significantly higher values in spring than in fall. We conclude that fine root biomass, especially in the finest fraction (< 1 mm in diameter), is strongly dependent on stand age. Among stands, carbon concentration in fine root biomass was highest in the 30-year-old stand, and appeared to be associated with the high tree and canopy density during the early stage of stand development. Values of RLD and RSA, morphological features indicative of stand nutrient-uptake efficiency, were higher in the 30-year-old stand than in the other stands.     

Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures

Since individual tree leaf area is an important measure for productivity as well as for site occupancy, it is of high interest in many studies about forest growth. The exact determination of leaf area is nearly impossible. Thus, a common way to get information about leaf area is to use substitutes. These substitutes are often variables which are collected in a destructive way which is not feasible for long term studies. Therefore, this study aimed at testing the applicability of using substitutes for leaf area which could be collected in a non-destructive way, namely crown surface area and crown projection area. In 8 stands of Norway spruce (Picea abies L. Karst.), divided into three age classes and two thinning treatments, a total of 156 trees were felled in order to test the relationship between leaf area and crown surface area and crown projection area, respectively. Individual tree leaf area of the felled sample trees was estimated by 3P-branch sampling with an accuracy of ±10%. Crown projection area and crown surface area were compared with other, more commonly used, but destructive predictors of leaf area, namely sapwood area at different heights on the bole. Our investigations confirmed findings of several studies that sapwood area is the most precise measure for leaf area because of the high correlation between sapwood area and the leaf area. But behind sapwood area at crown base and sapwood area at three tenth of the tree height the predictive ability of crown surface area was ranked third and even better than that of sapwood area at breast height (R² = 0.656 compared with 0.600). Within the stands leaf area is proportional to crown surface area. Using the pooled data of all stands a mixed model approach showed that additionally to crown surface area dominant height and diameter at breast height (dbh) improved the leaf area estimates. Thus, taking dominant height and dbh into account, crown surface area can be recommended for estimating the leaf area of individual trees. The resulting model was in line with many other findings on the leaf area and leaf mass relationships with crown size. From the additional influence of dominant height and dbh in the leaf area model we conclude that the used crown model could be improved by estimating the position of the maximum crown width and the crown width at the base of the crown depending on these two variables.     

Calibrating the liana crown occupancy index in Amazonian forests

Lianas hold an important, but understudied, role in forest dynamics, however they are difficult to measure and detailed liana measurements are time consuming. Many researchers have therefore used an ordinal scale index, the crown occupancy index (COI), to describe the liana load carried by trees. Here we assess the overall effectiveness, in terms of accuracy, precision, repeatability and efficiency, of the COI in tropical forests. We relate the COI to more detailed liana measurements at the individual tree-level and site-level, comparing sites with different levels of liana infestation. Our results show (1) that the COI accurately measures individual tree and plot level liana loads, indicated by the strong correlations between the COI and the number and basal area of lianas. However, (2) as expected, the COI is only weakly related to the basal area of lianas rooted close to the tree, which is a proxy for competition for below-ground resources. The COI is also (3) an efficient measure of liana loads, as the input time needed for a COI survey is considerably less than that of a detailed liana survey. We also (4) found a high degree of repeatability in COI classification between observers. Additionally (5), the COI can be used to differentiate between sites in terms of their overall liana canopy competition (precision), but (6) may not be a precise indicator of the site-level mean basal area of lianas in tree crowns.