Forest transpiration—targeted through xylem sap flux assessment versus hydrological modeling

The assessment of forest transpiration rates is crucial for determining plant-available soil water consumption and drought risk of trees. Xylem sap flux measurements have been used increasingly to quantify stand transpiration in forest ecosystems. Here, we compare this empirical approach with hydrological modeling on the basis of a stand transpiration dataset of adult beech (Fagus sylvatica), which was acquired across Bavaria, Germany, at eight forest sites. Xylem sap flux sensors were installed in five dominant trees each. Two tree to stand upscaling approaches, related to site-specific (1) sapwood area or (2) to leaf area index, were compared. The outcome was examined each in relation to process-based stand hydrological modeling, using LWF-BROOK90. Distinct relationships between tree diameter at breast height (1.30 m) and sapwood area-weighted sap flux along the radial profile became apparent across the study sites, confirming a generic allometric basis for stand-level upscaling of transpiration. The two upscaling approaches did not differ in outcome, representatively covering stand structure for comparison with modeling. Differential analysis yielded high agreement between the empirical and modeling approaches throughout most of the study period, although LWF-BROOK90 tended to overestimate sap flux measurements under low soil moisture. The two empirical approaches proved reliable for even-aged beech stands, as performance under high stand-structural heterogeneity awaits clarification. Findings advance stand-level hydrological modeling regarding coverage of stomatal behavior during temporary limitation in water availability.     

Why tree-crop interactions in agroforestry appear at odds with tree-grass interactions in tropical savannahs

This paper describes recent research findings on resource sharing between trees and crops in the semiarid tropics and attempts to reconcile this information with current knowledge of the interactions between savannah trees and understorey vegetation by examining agroforestry systems from the perspective of succession. In general, productivity of natural vegetation under savannah trees increases as rainfall decreases, while the opposite occurs in agroforestry. One explanation is that in the savannah, the beneficial effects of microclimatic improvements (e.g. lower temperatures and evaporation losses) are greater in more xeric environments. Mature savannah trees have a high proportion of woody above-ground structure compared to foliage, so that the amount of water 'saved' (largely by reduction in soil evaporation) is greater than water 'lost' through transpiration by trees. By contrast, in agroforestry practices such as alley cropping where tree density is high, any beneficial effects of the trees on microclimate are negated by reductions in soil moisture due to increasing interception losses and tree transpiration. While investment in woody structure can improve the water economy beneath agroforestry trees, it inevitably reduces the growth rate of the trees and thus increases the time required for improved understorey productivity. Therefore, agroforesters prefer trees with more direct and immediate benefits to farmers. The greatest opportunity for simultaneous agroforestry practices is therefore to fill niches within the landscape where resources are currently under-utilised by crops. In this way, agroforestry can mimic the large scale patch dynamics and successional progression of a natural ecosystem. This revised version was published online in June 2006 with corrections to the Cover Date.     

Estimating stand water use of large mountain ash trees and validation of the sap flow measurement technique

Mountain ash (Eucalyptus regnans F.J. Muell.) forest catchments exhibit a strong relationship between stand age and runoff, attributed inter alia to differences in tree water use. However, the tree water use component of the mountain ash forest water balance is poorly quantified. We have used the sap flow technique to obtain estimates of daily water use in large mountain ash trees. First, the sap flow technique was validated by means of an in situ cut tree experiment. Close agreement was obtained between the sap flow estimate of water use and the actual uptake of water by the tree from a reservoir. Second, we compared the variability in sap velocity between a symmetric and an asymmetric tree by using multiple sap flow loggers. In the symmetric tree, velocity was fairly uniform throughout the xylem during the day, indicating that accurate sap flow estimates can be obtained with a minimal number of sampling points. However, large variations in sap velocity were observed in the asymmetric tree, indicating that much larger sampling sizes are required in asymmetric stems for an accurate determination of mean sap velocity. Finally, we compared two procedures for scaling individual tree sap flow estimates to the stand level based on stem diameter and leaf area index measurements. The first procedure was based on a regression between stem diameter and tree water use, developed on a small sample of trees and applied to a stand-level census of stem diameter values. Inputs to the second procedure were tree water use and leaf area of a single tree and the leaf area index of the stand. The two procedures yielded similar results; however, the first procedure was more robust but it required more sampling effort than the second procedure.     

Using sap flow gauges to quantify water uptake by tree roots from beneath the crop rooting zone in agroforestry systems

Grevillea (Grevillea robusta A. Cunn.; Proteaceae) is used in agroforestry in many areas of the highlands of East and Central Africa, and is reported to be mainly deep rooted, with few shallow roots and correspondingly low levels of competition with associated crops for water and nutrients. To examine the extent of below-ground complementarily in water use between grevillea and cowpea (Vigna unguiculata L.; Leguminosae), experiments were carried out at the International Centre for Research in Agroforestry (ICRAF) Field Centre at Machakos, Kenya. Sap flux was measured using heat balance gauges attached to the stems of young grevillea trees (10–18 months old), both before and after excavating the crop rooting zone (upper 60 cm of soil) around the stem base, in order to establish the capacity of the grevillea to extract water from below this zone. After excavation, the trees maintained sap fluxes of up to 85% of the unexcavated values, suggesting a high degree of below-ground complementarity.     

Soil water dynamics in cropping systems containing <Emphasis Type="Italic">Gliricidia sepium</Emphasis>, pigeonpea and maize in southern Malawi

The water dynamics of cropping systems containing mixtures of Gliricidia sepium (Jacq.) Walp trees with maize (Zea mays L.) and/or pigeonpea (Cajanus cajan L.) were examined during three consecutive cropping seasons. The trees were pruned before and during each cropping season, but were left unpruned after harvesting the maize; prunings were returned to the cropping area in all agroforestry systems to provide green leaf manure. The hypothesis was that regular severe pruning of the trees would minimise competition with crops for soil moisture and enhance their growth by providing additional nutrients. Neutron probe measurements were used to determine spatial and temporal changes in soil moisture content during the 1997/98, 1998/99 and 1999/00 cropping seasons for various cropping systems. These included gliricidia intercropped with maize, with and without pigeonpea, a maize + pigeonpea intercrop, sole maize, sole pigeonpea and sole gliricidia. Soil water content was measured to a depth of 150 cm in all treatments at 4–6 week intervals during the main cropping season and less frequently at other times. Competition for water was apparently not a critical factor in determining crop performance as rainfall exceeded potential evaporation during the cropping season in all years. The distribution of water in the soil profile was generally comparable in all cropping systems, implying there was no spatial complementarity in water abstraction by tree and crop roots. However, available soil water content at the beginning of the cropping season was generally lower in the tree-based systems, suggesting that the trees continued to deplete available soil water during the dry season. The results show that, under rainfall conditions typical of southern Malawi, the soil profile contains sufficient stored water during the dry season (ca. 75–125 mm) to support the growth of gliricidia and pigeonpea, and that gliricidia trees pruned before and during the cropping season did not deleteriously compete for water with associated crops. Water use efficiency also appeared to be higher in the tree-based systems than in the sole maize and maize + pigeonpea treatments, subject to the proviso that the calculations were based on changes in soil water content rather than absolute measurements of water uptake by the trees and crops.     

Differences in transpiration between a forest and an agroforestry tree species in the Sudanian belt

Average population growth in the African Sudanian belt is 3 % per year. This leads to a significant increase in cultivated areas at the expense of fallows and forests. For centuries, rural populations have been practicing agroforestry dominated by Vitellaria paradoxa parklands. We wanted to know whether agroforestry can improve local rainfall recycling as well as forest. We compared transpiration and its seasonal variations between Vitellaria paradoxa, the dominant species in fallows, and Isoberlinia doka, the dominant species in dry forests in the Sudanian belt. The fallow and dry forest we studied are located in northwestern Benin, where average annual rainfall is 1200 mm. Sap flow density (SFD) was measured by transient thermal dissipation, from which tree transpiration was deduced. Transpiration of five trees per species was estimated by taking into account the radial profile of SFD. The effect of the species and of the season on transpiration was tested with a generalized linear mixed model. Over the three-year study period, daily transpiration of the agroforestry trees, V. paradoxa (diameters 8–38 cm) ranged between 4.4 and 26.8 L day^?1 while that of the forest trees, I. doka, (diameters 20–38 cm) ranged from 9.8 to 92.6 L day^?1. Daily transpiration of V. paradoxa was significantly lower (15 %) in the dry season than in the rainy season, whereas daily transpiration by I. doka was significantly higher (13 %) in the dry season than in the rainy season. Our results indicate that the woody cover of agroforestry systems is less efficient in recycling local rainfall than forest cover, not only due to lower tree density but also to species composition.     

Stored water use and transpiration in Scots pine: a modeling analysis with ANAFORE

We estimated daily use of stored water by Scots pine (Pinus sylvestris L.) trees growing in a temperate climate with the ANAFORE model (ANAlysis of FORest Ecosystems) and compared the simulation results with sap flow measurements. The original model was expanded with a dynamic water flow and storage model that simulates sap flow dynamics in an individual tree. ANAFORE was able to accurately simulate diurnal patterns of measured sap flow under microclimatic conditions that differ from those of the calibration period. Strong relationships were found between stored water use and several tree characteristics (diameter at breast height, sapwood area, leaf area), but not with tree height. Relative to transpiration, stored water use varied over time (between < 1% and 44% of daily transpiration). On days when transpiration was high, trees were more dependent on stored water, indicating that the contribution of internal water to transpiration is not a constant in the water budget of trees.     

Allometric estimation of above-ground biomass and leaf area in managed Grevillea robusta agroforestry systems

Non-destructive methods for determining the biomass and leaf area of individual trees throughout their growing cycle are an essential tool in agroforestry research, but must be capable of providing reliable estimates despite the influence that management strategies such as pruning may have on tree form. In the present study, allometric methods involving measurements of the diameter of all branches provided reliable estimates of canopy leaf area and biomass for grevillea trees (Grevillea robusta A. Cunn.; Proteaceae) grown as poles, but proved unsuitable for routine measurements because of their time-consuming nature. An alternative, less laborious method based on measurements of trunk cross-sectional area immediately below the first branch of the canopy provided satisfactory allometric estimates of leaf area and canopy biomass. Trunk biomass was determined from measurements of tree height and diameter at breast height using established methodology based on the assumption that trunk volume may be calculated using a quadratic paraboloid model; biomass was determined as the product of trunk volume and the specific gravity of the wood. The theoretical basis, development and validation of allometric methods for estimating tree growth are discussed and their wider applicability to other agroforestry systems is assessed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Tree water use and rainfall partitioning in a mature poplar-pasture system

Traditionally, poplars (Populus) have been planted to control erosion on New Zealand's hill-slopes, because of their capacity to dry out and bind together the soil, by reducing effective rainfall and increasing evapotranspiration and soil strength. However, the effect of widely spaced poplars on the partitioning of soil water and rainfall has not been reported. This study determined rainfall partitioning for 18 mid-spring days in a mature P. deltoides (Bart. ex Marsh, Clone I78)-pasture association (37 stems per hectare, unevenly spaced at 16.4 +/- 0.4 m) and compared it with a traditional open pasture system in grazed areas of a hill environment. Tree transpiration was measured by the heat pulse technique. A time-driven mathematical model was used to set a zero offset, adjust anomalous values and describe simultaneous sap velocity time courses of trees. The model showed that daylight sap flow velocities can be represented with a nonlinear Beta function (R(2) > 0.98), and differences in the parameters representing the initiation, duration and conformation of the sap velocity can be tested statistically to discern tree transpiration differences during the day. Evapotranspiration was greater for the poplar-pasture association than for the open pasture (2.7-3.0 versus 2.2 mm day(-1)). The tree canopy alone contributed 0.92 mm day(-1) as transpiration and 1.37 mm day(-1) as interception, whereas evapotranspiration of the pasture understory was only 0.4-0.6 mm day(-1). Despite the higher water use of the poplar-pasture association, soil water in the 0-300 mm soil stratum was higher than, or similar to, that of the open pasture. Tree shading decreased evapotranspiration and pasture accumulation under the trees.     

Soil water competition in a temperate hedgerow agroforestry system in South Africa

An on-farm trial was conducted to determine dry matter production of four fodder tree species and their effect on soil water and maize production. The trees were planted in rows intercropped with maize. The four tree species selected were Acacia karroo Hayne (indigenous fodder tree), Leucaena leucocephala (Lam.) De Wit (nitrogen fixing), Morus alba L. (fodder and fruit), and Gleditsia triacanthos L. (fodder and fuel). Volumetric soil water was measured in the upper 0.3 m of soil in each row of the trial using the time domain reflectometry technique. The neutron probe technique was used for monitoring the water content deeper in the soil. Geostatistical methods were used to analyse treatment differences in the upper 0.3 m of soil. The soil water content did not differ significantly between the maize and tree rows indicating that competition for water in the upper horizon was not the reason for lower maize yields. However, at greater soil depths (75–125 cm) trees in the wide spacing used less water than those in the narrow spacing. Light interception was an important factor in reducing maize yields in the row nearest to the trees. High soil water values recorded during summer indicated that in the current cycle of good rainfall the plants in the agroforestry trial were not stressed. Thus the trees do not compete with the crops for soil moisture in good rainfall seasons. However, this study would need further evaluation for the competition for water for the low rainfall years. Since the trees have access to water at greater depths, they are likely to be more productive into the dry season than shallow rooted crops.     

Effects of thinning on soil and tree water relations, transpiration and growth in an oak forest (Quercus petraea (Matt.) Liebl.)

To quantify the effects of crown thinning on the water balance and growth of the stand and to analyze the ecophysiological modifications induced by canopy opening on individual tree water relations, we conducted a thinning experiment in a 43-year-old Quercus petraea stand by removing trees from the upper canopy level. Soil water content, rainfall interception, sap flow, leaf water potential and stomatal conductance were monitored for two seasons following thinning. Seasonal time courses of leaf area index (LAI) and girth increment were also measured. Predawn leaf water potential was significantly higher in trees in the thinned stand than in the closed stand, as a consequence of higher relative extractable water in the soil. The improvement in water availability in the thinned stand resulted from decreases in both interception and transpiration. From Year 1 to Year 2, an increase in transpiration was observed in the thinned stand without any modification in LAI, whereas changes in transpiration in the closed stand were accompanied by variations in LAI. The different behaviors of the closed and open canopies were interpreted in terms of coupling to the atmosphere. Thinning increased inter-tree variability in sap flow density, which was closely related to a leaf area competition index. Stomatal conductance varied little inside the crown and differences in stomatal conductance between the treatments appeared only during a water shortage and affected mainly the closed stand. Thinning enhanced tree growth as a result of a longer growing period due to the absence of summer drought and higher rates of growth. Suppressed and dominant trees benefited more from thinning than trees in the codominant classes.     

Structural and compositional controls on transpiration in 40- and 450-year-old riparian forests in western Oregon, USA

Large areas of forests in the Pacific Northwest are being transformed to younger forests, yet little is known about the impact this may have on hydrological cycles. Previous work suggests that old trees use less water per unit leaf area or sapwood area than young mature trees of the same species in similar environments. Do old forests, therefore, use less water than young mature forests in similar environments, or are there other structural or compositional components in the forests that compensate for tree-level differences? We investigated the impacts of tree age, species composition and sapwood basal area on stand-level transpiration in adjacent watersheds at the H.J. Andrews Forest in the western Cascades of Oregon, one containing a young, mature (about 40 years since disturbance) conifer forest and the other an old growth (about 450 years since disturbance) forest. Sap flow measurements were used to evaluate the degree to which differences in age and species composition affect water use. Stand sapwood basal area was evaluated based on a vegetation survey for species, basal area and sapwood basal area in the riparian area of two watersheds. A simple scaling exercise derived from estimated differences in water use as a result of differences in age, species composition and stand sapwood area was used to estimate transpiration from late June through October within the entire riparian area of these watersheds. Transpiration was higher in the young stand because of greater sap flux density (sap flow per unit sapwood area) by age class and species, and greater total stand sapwood area. During the measurement period, mean daily sap flux density was 2.30 times higher in young compared with old Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees. Sap flux density was 1.41 times higher in young red alder (Alnus rubra Bong.) compared with young P. menziesii trees, and was 1.45 times higher in old P. menziesii compared with old western hemlock (Tsuga heterophylla (Raf.) Sarg.) trees. Overall, sapwood basal area was 21% higher in the young stand than in the old stand. In the old forest, T. heterophylla is an important co-dominant, accounting for 58% of total sapwood basal area, whereas P. menziesii is the only dominant conifer in the young stand. Angiosperms accounted for 36% of total sapwood basal area in the young stand, but only 7% in the old stand. For all factors combined, we estimated 3.27 times more water use by vegetation in the riparian area of the young stand over the measurement period. Tree age had the greatest effect on stand differences in water use, followed by differences in sapwood basal area, and finally species composition. The large differences in transpiration provide further evidence that forest management alters site water balance via elevated transpiration in vigorous young stands.     

Exploring below ground complementarity in agroforestry using sap flow and root fractal techniques

Indices of shallow rootedness and fractal methods of root system study were combined with sapflow monitoring to determine whether these ‘short-cut’ methods could be used to predict tree competition with crops and complementarity of below ground resource use in an agroforestry trial in semi-arid Kenya. These methods were applied to Grevillea robusta Cunn., Gliricidia sepium (Jacq.) Walp., Melia volkensii Gürke and Senna spectabilis syn. Cassia spectabilis aged two and four years which were grown in simultaneous linear agroforestry plots with maize as the crop species. Indices of competition (shallow rootedness) differed substantially according to tree age and did not accurately predict tree:crop competition in plots containing trees aged four years. Predicted competition by trees on crops was improved by multiplying the sum of proximal diameters squared for shallow roots by diameter at breast height², thus taking tree size into account. Fractal methods for the quantification of total length of tree root systems worked well with the permanent structural root system of trees but seriously underestimated the length of fine roots (less than 2 mm diameter). Sap flow measurements of individual roots showed that as expected, deep tap roots provided most of the water used by the trees during the dry season. Following rainfall, substantial water uptake by shallow lateral roots occurred more or less immediately, suggesting that existing roots were functioning in the recently wetted soil and that there was no need for new fine roots to be produced to enable water uptake following rainfall. This revised version was published online in June 2006 with corrections to the Cover Date.     

Estimating sap flux densities in date palm trees using the heat dissipation method and weighing lysimeters

In a world of diminishing water reservoirs and a rising demand for food, the practice and development of water stress indicators and sensors are in rapid progress. The heat dissipation method, originally established by Granier, is herein applied and modified to enable sap flow measurements in date palm trees in the southern Arava desert of Israel. A long and tough sensor was constructed to withstand insertion into the date palm's hard exterior stem. This stem is wide and fibrous, surrounded by an even tougher external non-conducting layer of dead leaf bases. Furthermore, being a monocot species, water flow does not necessarily occur through the outer part of the palm's stem, as in most trees. Therefore, it is highly important to investigate the variations of the sap flux densities and determine the preferable location for sap flow sensing within the stem. Once installed into fully grown date palm trees stationed on weighing lysimeters, sap flow as measured by the modified sensors was compared with the actual transpiration. Sap flow was found to be well correlated with transpiration, especially when using a recent calibration equation rather than the original Granier equation. Furthermore, inducing the axial variability of the sap flux densities was found to be highly important for accurate assessments of transpiration by sap flow measurements. The sensors indicated no transpiration at night, a high increase of transpiration from 06:00 to 09:00, maximum transpiration at 12:00, followed by a moderate reduction until 08:00; when transpiration ceased. These results were reinforced by the lysimeters' output. Reduced sap flux densities were detected at the stem's mantle when compared with its center. These results were reinforced by mechanistic measurements of the stem's specific hydraulic conductivity. Variance on the vertical axis was also observed, indicating an accelerated flow towards the upper parts of the tree and raising a hypothesis concerning dehydrating mechanisms of the date palm tree. Finally, the sensors indicated reduction in flow almost immediately after irrigation of field-grown trees was withheld, at a time when no climatic or phenological conditions could have led to reduction in transpiration.     

Tree/crop complementarity in an arid zone runoff agroforestry system in northern Kenya

We tested the hypothesis that shallow-rooted crops and deep-rooted trees will share the available water in a complementary manner, when grown together, in a field trail in the Turkana district of northern Kenya during 1994 to 1996. Such studies have been few in dryland agroforestry. The effects of two different Acacia saligna (Labill.) H. Wendl. tree planting densities (2500 and 833 trees per ha), tree pruning (no pruning vs. pruning) and annual intercrops (no intercrop vs. intercrop) on total biomass production and their interactions were tested. In 1996 Sorghum bicolor (L.) Moench was used during the first vegetation period and Vigna unguiculata (L.) Walp. during the second. We used naturally generated runoff water for irrigation to supplement low rainfall amounts typical for the area. High biomass production (> 13 t ha^–1 over a two year period) was observed irrespective of intercropping of pruned trees or sole tree stands. Although the pruning treatment reduced total tree biomass yields by a quarter, the introduction of annual intercrops after the pruning of trees outweighed this loss. The yields of the intercrops in the pruned tree treatments were similar to their yields when grown as monocrops. The calculation of land equivalent ratios showed overyielding for intercropped, pruned systems. The high values for LER (1.36 at low and 1.47 at high density of trees) indicate that there is complementarity in resource use between the different species.This revised version was published online in November 2005 with corrections to the Cover Date.     

Linking leaf and tree water use with an individual-tree model

We tested the ability of a model to scale gas exchange from leaf level to whole-tree level by: (1) measuring leaf gas exchange in the canopy of 10 trees in a tall Eucalyptus delegatensis RT Baker forest in NSW, Australia; (2) monitoring sap flow of the same 10 trees during the measurement week; and (3) using an individual-tree-based model (MAESTRA) to link the two sets of measurements. Photosynthesis and stomatal conductance components of the model were parameterized with the leaf gas exchange data, and canopy structure was parameterized with crown heights, dimensions and leaf areas of each of the measurement trees and up to 45 neighboring trees. Transpiration of the measurement trees was predicted by the model and compared with sap flow data. Leaf gas exchange parameters were similar for all 10 trees, with the exception of two smaller trees that had relatively low stomatal conductances. We hypothesize that these trees may have experienced water stress as a result of competition from large neighboring trees. The model performed well, and in most cases, was able to replicate the time course of tree transpiration. Maximum rates of transpiration were higher than measured rates for some trees and lower than measured rates for others, which may have been a result of inaccuracy in estimating tree leaf area. There was a small lag (about 15-30 minutes) between sap flow and modeled transpiration for some trees in the morning, likely associated with use of water stored in stems. The model also captured patterns of variation in sap flow among trees. Overall, the study confirms the ability of models to estimate forest canopy transpiration from leaf-level measurements.     

Dynamics of transpiration, sap flow and use of stored water in tropical forest canopy trees

In large trees, the daily onset of transpiration causes water to be withdrawn from internal storage compartments, resulting in lags between changes in transpiration and sap flow at the base of the tree. We measured time courses of sap flow, hydraulic resistance, plant water potential and stomatal resistance in co-occurring tropical forest canopy trees with trunk diameters ranging from 0.34-0.98 m, to determine how total daily water use and daily reliance on stored water scaled with size. We also examined the effects of scale and tree hydraulic properties on apparent time constants for changes in transpiration and water flow in response to fluctuating environmental variables. Time constants for water movement were estimated from whole-tree hydraulic resistance (R) and capacitance (C) using an electric circuit analogy, and from rates of change in water movement through intact trees. Total daily water use and reliance on stored water were strongly correlated with trunk diameter, independent of species. Although total daily withdrawal of water from internal storage increased with tree size, its relative contribution to the daily water budget (approximately 10%) remained constant. Net withdrawal of water from storage ceased when upper branch water potential corresponded to the sapwood water potential (Psi(sw)) at which further withdrawal of water from sapwood would have caused Psi(sw) to decline precipitously. Stomatal coordination of vapor and liquid phase resistances played a key role in limiting stored water use to a nearly constant fraction of total daily water use. Time constants for changes in transpiration, estimated as the product of whole- tree R and C, were similar among individuals (~0.53 h), indicating that R and C co-varied with tree size in an inverse manner. Similarly, time constants estimated from rates of change in crown and basal sap flux were nearly identical among individuals and therefore independent of tree size and species.     

Spatial distribution and sampling of Thaumetopoea pityocampa (Den. & Schiff.) (Lep. Thaumetopoeidea) populations on Pinus pinaster Ait. in Montesinho, N. Portugal

The pine processionary moth, Thaumetopoea pityocampa (Den. & Schiff.) is one of the most destructive pine pests in the Mediterranean countries. The first objective of this study was to analyse the spatial distribution of this insect on the tree on Pinus pinaster Ait., in northeast Portugal. The second objective was to develop two sampling plans: a sampling plan with fixed levels of precision for estimating T. pityocampa populations and a sequential sampling plan to classify the pest. Population estimates were made by registering the number of insect egg batches on 90 trees distributed over 3 stands during a 3-year study (1997–2000). Taylor's power law and Iwao regression were used to analyse the spatial distribution of the pest. Taylor's power law fitted the data better (r² = 0.775) than Iwao's regression model (r² = 0.704). The aggregation indices (b and β) were both significantly greater than one, indicating the aggregation of T. pityocampa egg batches. Optimal sample sizes for fixed precision levels of 0.10, 0.15 and 0.25 were estimated with Taylor's regression coefficients. Required sample sizes increased dramatically with increased levels of precision. The two sampling plans presented should be a tool for research on population dynamics and pest management decision.     

Estimating the water use of a sclerophyllous species under an East-Mediterranean climate: I. Response of transpiration of Phillyrea latifolia L. to site factors

Water use by vegetation is the most important parameter which is directly related to plant anatomical and eco-physiological characters, and to soil–bedrock water availability. This basic parameter is important in decision making, planning and management of the open areas under semiarid climatic conditions like those of Israel. Attempts were made to quantify the water use by Phillyrea latifolia L. by means of the heat pulse method. This method provides continuous monitoring of the convective heat pulse velocity (HPV) in the trunk xylem, thus facilitating the quantification of the transpiration flux per tree. The daily and annual water use by this species was estimated by extrapolations from the single-tree transpiration rates, based on the relationship between the leaf area of the measured trees and the entire area.

The lithological properties of the bedrock formations in each of the research sites caused significant differences between sites in the average daily transpiration rate and, hence, in the total water use by the P. latifolia scrub formation. The daily transpiration, averaged over a year, ranged between 4.00 and 8.15 l per day per tree, and the estimated annual water use ranged between 179.4 and 365.5 mm, i.e., between 30 and 61% of the annual rainfall in the winter preceding the measurements in 1993.     

Estimating water use of sclerophyllous species under East-Mediterranean climate: III. Tabor oak forest sap flow distribution and transpiration

The annual course of daily transpiration and the hydrological balance of a Tabor oak forest were determined. The study was done in a representative forest within the natural geographical range of the species in the lower Galilee region of Israel. The climate is sub-humid with a rainless dry season from May to October. A partial water balance of a 0.1 ha area supporting an average of 14 trees was calculated from: (a) soil water content (SWC) measured by a Neutron Probe at depths of from 0.2 to 8 m, and (b) daylight transpiration rate measured with sap flow sensors by the heat pulse technique.Soil–bedrock complex water content (%) in the first 2 m of the profile fluctuated strongly between 5 and 20% depending on the season. The water content increased with depth from about 10% at 2.0 m depth to more than 20% at 5.0 m depth. For depths exceeding 5.5 m seasonal fluctuations in water content were negligible and water content ranged from 30 to 35%. After a dry winter, water content generally decreased within the main root zone down to about 2.0 m depth. Monthly changes in water content (mm) were greatest at depths of 0.35–1.0 m. Only minor changes in the soil–bedrock complex water content were recorded at greater depths. After a very rainy winter (2002/2003), decreases in soil–bedrock complex water content in the upper 2 m were much larger than after a dry winter. Fluctuations of soil–bedrock complex water content in deeper regions were larger in the wetter year, probably the result of drainage.Sap velocity was measured at six depths in the sapwood, from 4 to 44 mm, at 8 mm intervals. Sap velocity declined with depth, hence, sap flux density too.Based on sap velocity measurements performed during 4 years, the annual average daily transpiration (T) was 0.796 mm/day. This sums up to 239 mm during ∼300 days of leaf carriage, i.e. 41.3% of the 578 mm average annual rainfall for the area in the last 50 years. In a relatively dry year (rainfall of 432.7 mm) total water withdrawal from the 8 m soil–bedrock profile was 81% of the annual rainfall; of this amount 69% were transpired by the oak trees (239.0 mm), or 55% of the annual rainfall. In a relatively wet year (annual rainfall 801.4 mm) total water withdrawal was 67%; of this amount 45% would be transpired by the oak trees, or 30% of the annual rainfall.