Abstract: Although the global partitioning of evapotranspiration (ET) into transpiration, soil evaporation, and canopy evaporation is not well known, most current land surface schemes and the few available observations indicate that transpiration is the dominant component on the global scale, followed by soil evaporation and canopy evaporation. The Community Land Model version 3 (CLM3), however, does not reflect this global view of ET partitioning, with soil evaporation and canopy evaporation far outweighing transpiration. One consequence of this unrealistic ET partitioning in CLM3 is that photosynthesis, which is linked to transpiration through stomatal conductance, is significantly underestimated on a global basis. A number of modifications to CLM3 vegetation and soil hydrology parameterizations are described that improve ET partitioning and reduce an apparent dry soil bias in CLM3. The modifications reduce canopy interception and evaporation, reduce soil moisture stress on transpiration, increase transp...

Abstract: [1] The shear layer at the top of a submerged canopy generates coherent vortices that control exchange between the canopy and the overflowing water. Unlike free shear layers, the vortices in a canopy shear layer do not grow continuously downstream but reach and maintain a finite scale determined by a balance between shear production and canopy dissipation. This balance defines the length scale of vortex penetration into the canopy, δe, and the region of rapid exchange between the canopy and overflow. Deeper within the canopy, transport is constrained by smaller turbulence scales. A two-box canopy model is proposed on the basis of the length scale δe. Using diffusivity and exchange rates defined in previous studies, the model predicts the timescale required to flush the canopy through vertical exchange over a range of canopy density and height. The predicted canopy retention times, which range from minutes to an hour, are consistent with canopy retention inferred from tracer observations in the field and comparable to retention times for some hyporheic regions. The timescale for vertical exchange, along with the in-canopy velocity, determines the minimum canopy length for which vertical exchange dominates water renewal. Shorter canopies renew interior water through longitudinal advection. Finally, canopy water retention influences longitudinal dispersion through a transient storage process. When vertical exchange controls canopy retention, the transient storage dispersion increases with canopy height. When longitudinal advection controls water renewal, dispersion increases with canopy patch length.

Abstract: Stomatal responses to atmospheric change have been well documented through a range of laboratory- and field-based experiments. Increases in atmospheric concentration of CO2 ([CO2]) have been shown to decrease stomatal conductance (gs) for a wide range of species under numerous conditions. Less well understood, however, is the extent to which leaf-level responses translate to changes in ecosystem evapotranspiration (ET). Since many changes at the soil, plant, and canopy microclimate levels may feed back on ET, it is not certain that a decrease in gs will decrease ET in rain-fed crops. To examine the scaling of the effect of elevated [CO2] on gs at the leaf to ecosystem ET, soybean (Glycine max) was grown in field conditions under control (approximately 375 μmol CO2 mol−1 air) and elevated [CO2] (approximately 550 μmol mol−1) using free air CO2 enrichment. ET was determined from the time of canopy closure to crop senescence using a residual energy balance approach over four growing seasons. Elevated [CO2] caused ET to decrease between 9% and 16% depending on year and despite large increases in photosynthesis and seed yield. Ecosystem ET was linked with gs of the upper canopy leaves when averaged across the growing seasons, such that a 10% decrease in gs results in a 8.6% decrease in ET; this relationship was not altered by growth at elevated [CO2]. The findings are consistent with model and historical analyses that suggest that, despite system feedbacks, decreased gs of upper canopy leaves at elevated [CO2] results in decreased transfer of water vapor to the atmosphere.

Abstract: For three forest canopies (a sparse, boreal needleleaf; a temperate broadleaf; and a dense, tropical, broadleaf stand) light-use efficiency (LUE) is found to be 6-33% higher when sky radiance is dominated by diffuse rather than direct sunlight. This enhancement is much less than that reported previously for both crops (110%; Choudbury, 2001 ) and moderately dense temperate woodland (50-180%). We use the land-surface scheme JULES to interpret the observed canopy response. Once sunflecks and leaf orientation are incorporated explicitly into the scheme, our simulations reproduce convincingly the overall level of canopy gross photosynthetic product (GPP), its enhancement with respect to diffuse sunlight and the mean 15% reduction in productivity observed during the afternoon due to stomatal closure. The LUE enhancement under diffuse sunlight can be explained by sharing of the canopy radiation-load, which is reduced under direct sky radiance. Once sunflecks are accounted for the advantage of implementing more sophisticated calculations of stomatal conductance (e.g. Ball-Berry and SPA submodels) is less obvious even for afternoon assimilation. Empirical relations are developed between observed carbon flux and the environmental variables total downwelling shortwave radiation (SW), canopy temperature (T) and the fraction of diffuse sky radiance (fDIF). These relations allow us to gauge the impact of increased/reduced insolation on GPP and net ecosystem exchange (NEE). Overall the three stands appear to be fairly stable within global trends and typical interannual variability (SW changing by <15%). Greatest sensitivity is exhibited by the boreal site, Zotino, where NEE falls by 9±4% for a 15% reduction in SW.

Abstract: Cloud cover increases the proportion of diffuse radiation reaching the Earth’s surface and affects many microclimatic factors such as temperature, vapour pressure deficit and precipitation. We compared the relative efficiencies of canopy photosynthesis to diffuse and direct photosynthetic photon flux density (PPFD) for a Norway spruce forest (25year-old, leaf area index 11 m 2 m � 2 ) during two successive 7-day periods in August. The comparison was based on the response of net ecosystem exchange (NEE) of CO2 to PPFD. NEE and stomatal conductance at the canopy level (Gcanopy) was estimated from halfhourly eddy-covariance measurements of CO2 and H2O fluxes. In addition, daily courses of CO2 assimilation rate (AN) and stomatal conductance (Gs) at shoot level were measured using a gas-exchange technique applied to branches of trees. The extent of spectral changes in incident solar radiation was assessed using a spectroradiometer. We found significantly higher NEE (up to 150%) during the cloudy periods compared with the sunny periods at corresponding PPFDs. Prevailing diffuse radiation under the cloudy days resulted in a significantly lower compensation irradiance (by ca. 50% and 70%), while apparent quantum yield was slightly higher (by ca. 7%) at canopy level and significantly higher (by ca. 530%) in sun-acclimated shoots. The main reasons for these differences appear to be (1) more favourable microclimatic conditions during cloudy periods, (2) stimulation of photochemical reactions and stomatal opening via an increase of blue/red light ratio, and (3) increased penetration of light into the canopy and thus a more equitable distribution of light between leaves. Our analyses identified the most important reason of enhanced NEE under cloudy sky conditions to be the effective penetration of diffuse radiation to lower depths of the canopy. This subsequently led to the significantly higher solar equivalent leaf area compared with the direct radiation. Most of the leaves in such dense canopy are in deep shade, with marginal or negative carbon balances during sunny days. These findings show that the energy of diffuse, compared with direct, solar radiation is used more efficiently in assimilation processes at both leaf and canopy levels.

Abstract: Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm2 g−1 whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.

Abstract: Thefluxcontributionofcoherentstructurestothetotalexchangeofenergy and matter is investigated in a spruce canopy of moderate density in heterogeneous, complex terrain. The study deploys two methods of analysis to estimate the coherent exchange: conditional averages in combination with wavelet analysis, and quadrant analysis. The data were obtained by high-frequency single-point measurements using sonic anemometers and gas analysers at five observation heights above and within the canopy and subcanopy, and represent a period of up to 2.5months. The study mainly addresses the momentum transfer and exchange of sensible heat throughout the roughness sublayer, while results are provided for the exchange of carbon dioxide and water vapour above the canopy. The magnitude of the flux contribution of coherent structures largely depends on the method of analysis, and it is demonstrated that these differences are attributed to differences in the sampling strategy between the two methods. Despite the differ- ences, relational properties such as sweep and ejection ratios and the variation of the flux contribution with height were in agreement for both methods. The sweep phase of coherentstructures is the dominantprocess close to and within the canopy, whereas the ejections gain importance with increasing distance to the canopy. The efficiency of the coherent exchange in transporting scalars exceeds that for momentum by a factor of two. The occurrence of coherent structures results in a flux error less than 4% for the eddy-covariance method. Based on the physical processes identified from the analysis of the ejection and sweep phases along the vertical profile in the rough- ness sublayer, a classification scheme for the identification of exchange regimes is developed. This scheme allows one to estimate the region of the canopy participat- ing in the exchange of energy and matter with the above-canopy air under varying environmental conditions.

Abstract: We studied species richness, composition, and vertical distribution of vascular epiphytes at two sites in the Bolivian Andes. To account for the epiphyte flora on understory trees, epiphytes on shrubs and small trees were sampled in 20 × 20 m2 subplots around each sampled canopy tree; this understory zone U is introduced as an addition to the well-established five vertical Johansson tree zones. More than 20% of about 500 species recorded were found only in the understory subplots, including ca. 40% of aroids, 35%–40% of piperoids, and 25%–30% of ferns. Habitat generalists (occurring in three or more zones) were most common, contributing about 50% of all species, specialists (occurring only in two zones, or in three continuous ones) 34%–42%, and hemiepiphytes 6%–16%. Canopy epiphytes (occurring > 90% in tree zones Z3–5) were mainly represented by orchids and ferns, many with special adaptations to drought stress such as pseudobulbs, succulence, and poikilohydry. Trunk epiphytes ( > 90% in understory and tree zones Z1–2) reached highest relative species numbers among piperoids and ferns. Most hemiepiphytes were also trunk epiphytes, due to their characteristic growth pattern, and included mainly aroids. The vertical distribution of epiphytes within a tree is determined by several microenvironmental gradients, with light intensity, wind speed, and air temperature increasing and air humidity decreasing from the ground level to the canopy.

Abstract: Background. 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. Methodology/Principal Findings. 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. Conclusions/Significance. 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.

Abstract: Leaf Area Index (leaf area per unit ground area, LAI) is a key driver of forest productivity but has never previously been measured directly at the landscape scale in tropical rain forest (TRF) We used a modular tower and stratified random sampling to harvest all foliage from forest floor to canopy top in 55 vertical transects (46 m 2 ) across 500 ha of old growth in Costa Rica Landscape LAI was 600 ± 032 SEM Trees, palms and lianas accounted for 89% of the total, and trees and lianas were 95% of the upper canopy All vertical transects were organized into quantitatively defined strata, partially resolving the long-standing controversy over canopy stratification in TRF Total LAI was strongly correlated with forest height up to 21 m, while the number of canopy strata increased with forest height across the full height range These data are a benchmark for understanding the structure and functional composition of TRF canopies at landscape scales, and also provide insights for improving ecosystem models and remote sensing validation

Abstract: Irrigation scheduling is critical as it affects both fruit yield and composition. We examined the potential to use field-measured hyperspectral remote sensing data (reflectance and transmission over the 350–2500 nm wavelength region) to estimate leaf water content, equivalent water thickness (EWT), and leaf water potential (Ψ) in a commercial vineyard of Vitis vinifera cv. Pinot noir. The data allowed us to evaluate a number of reflectance patterns to estimate vine water status through correlations using two spectral approaches: direct measurement of vegetation indexes (VIs) and continuum removal analysis (CRA). Continuum removal analysis was applied to obtain the maximum band depth (MBD) and the band area (BA) of several absorption features sensitive to water content. Correlations were high for EWT at the leaf level using a modification of the Simple Ratio VI (SR2; R2 = 0.916) and for CRA with MBD970 (R2 = 0.917) and BA1160 (R2 = 0.897). Correlations with EWT and water potential at the canopy level for SR2 were nonsignificant, which was characteristic for many VIs. For predawn water potential (ΨPD) and midday stem water potential (Ψstem) at the canopy level, best fits were realized for Modified Triangular VI (MTVI2; R2 = 0.360) and Red/Green VI (RGI, R695/R554; R2 = 0.462), respectively. For canopy level water status, the best results were obtained using the difference between the midday stem water potential and the pre-dawn leaf water potential (Ψstem - ΨPD) with R2 = 0.619 for RGI and R2 = 0.541 for Structure Intensive Pigment Index (SIPI, R800-R445/R800-R680), while for CRA R2 = 0.477 for BA1600 and R2 = 0.509 for MBD970. Results suggest that noninvasive monitoring using hyperspectral data could improve current methods for estimating water status in individual vines. Applications of similar measurements could be produced from airborne hyperspectral imagers to provide spatially resolved estimates of water stress for use in water management of large-scale commercial vineyards.

Abstract: The paper summarizes research conducted at International Center for Tropical Agriculture (CIAT) on responses of cassava to extended water shortages in the field aided by modern gas-exchange and water-relation techniques as well as biochemical assays. The aim of the research was to coordinate basic and applied aspects of crop physiology into a breeding strategy with a multidisciplinary approach. Several physiological characteristics/traits and mechanisms underpinning tolerance of cassava to drought were elucidated using a large number of genotypes from the CIAT core germplasm collection grown in various locations representing ecozones where cassava is cultivated. Most notable among these characteristics are the high photosynthetic capacity of cassava leaves in favorable environments and the maintenance of reasonable rates throughout prolonged water deficits, a crucial characteristic for high and sustainable productivity. Cassava possess a tight stomatal control over leaf gas exchange that reduces water losses when plants are subjected to soil water deficits as well as to high atmospheric evaporative demands, thus protecting leaves from severe dehydration. During prolonged water deficits, cassava reduces its canopy by shedding older leaves and forming smaller new leaves leading to less light interception, another adaptive trait to drought. Though root yield is reduced (but much less than the reduction in top growth) under water stress, the crop can recover when water becomes available by rapidly forming new canopy leaves with much higher photosynthetic rates compared to unstressed crops, thus compensating for yield losses with final yields approaching those in well-watered crops. Cassava can extract slowly water from deep soils, a characteristic of paramount importance in seasonally dry and semiarid environments where deeply stored water needs to be tapped. Screening large accessions under seasonally dry and semiarid environments showed that yield is significantly correlated with upper canopy leaf photosynthetic rates, and the association was attributed mainly to nonstomatal (anatomical/biochemical) factors. Parental materials with both high yields and photosynthetic rates were identified for incorporation into breeding and selection programs for cultivars adapted to prolonged drought coupled with high temperatures and dry air, conditions that might be further aggravated by global climate changes in tropical regions.

Abstract: The viticultural effects of sward competition with vines for water and nutrients, indirectly resulting from reduced herbicide use under a sustainable production system, are well established in cooler, more humid areas but are less well understood in warm and dry areas. Three groundcover treatments—standard industry practice of herbicide-treated undervine strips, bare soil maintained with herbicides, and complete floor cover (no herbicides)—were applied in two Chardonnay vineyards, one in a hot dry climate, the other in a mild climate. Increasing floor cover decreased early-season soil moisture and petiole nutrient status and strongly reduced vine vegetative growth. After three experimental years, grape yield also became limited. Large treatment differences in vegetative growth, canopy structure, and yield contributed to differences in berry weight and composition. Vine response to floor covers was less pronounced under mild climatic conditions compared to hot and dry conditions. Total or partial floor cover is a powerful tool for controlling vegetative growth of grapevines. However, under hot and dry conditions, competition for water and nutrients, particularly if it occurs at sensitive stages (such as bloom and berry set) can lead to a substantial decrease in yield and vine capacity.

Abstract: The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.

Abstract: Epiphytic biomass, canopy humus and associated canopy water storage capacity are known to vary greatly between old-growth tropical montane cloud forests but for regenerating forests such data are virtually absent. The present study was conducted in an old-growth cloud forest and in a 30-year-old secondary forest (SF) on wind-exposed slopes in the Cordillera de Tilaran (Monteverde area) in northern Costa Rica. Epiphytic vegetation in both forests was dominated by bryophytes. Epiphyte mat weight (epiphyte biomass and canopy humus) at the stand level was 1,035 kg ha−1 in the SF and 16,215 kg ha−1 in the old-growth forest (OGF). The water contents of epiphytic bryophytes in the OGF were determined gravimetrically in situ and showed maximum values of 418% ± 74 (SD)% of dry weight and minimum values of 36% ± 10 (SD)%. Maximum stand water storage of non-vascular epiphytes and canopy humus at Monteverde was estimated at 0.36 mm for the SF and 4.95 mm for the OGF. Epiphytic bryophytes exhibited more dynamic wetting and drying cycles compared to canopy humus. Maximum water loss through evaporation was 251% of dry weight (bryophytes) and 117% of dry weight (canopy humus) within 3 days of sunny weather without precipitation. Despite the high potential water storage capacity of epiphytic bryophytes and canopy humus the actually available storage is likely to be much smaller depending on antecedent rainfall and evaporative conditions.

Abstract: Increased canopy leaf area (L) may lead to higher forest productivity and alter processes such as species dynamics and ecosystem mass and energy fluxes. Few CO2 enrichment studies have been conducted in closed canopy forests and none have shown a sustained enhancement of L. We reconstructed 8 years (1996‐2003) of L at Duke’s Free Air CO2 Enrichment experiment to determine the effects of elevated atmospheric CO2 concentration ([CO2]) on L before and after canopy closure in a pine forest with a hardwood component, focusing on interactions with temporal variation in water availability and spatial variation in nitrogen (N) supply. The dynamics of L were reconstructed using data on leaf litterfall mass and specific leaf area for hardwoods, and needle litterfall mass and specific leaf area combined with needle elongation rates, and fascicle and shoot counts for pines. The dynamics of pine L production and senescence were unaffected by elevated [CO2], although L senescence for hardwoods was slowed. Elevated [CO2] enhanced pine L and the total canopy L (combined pine and hardwood species; Po0.050); on average, enhancement following canopy closure was � 16% and 14% respectively. However, variation in pine L and its response to elevated [CO2] was not random. Each year pine L under ambient and elevated [CO2 ]w asspatially correlated to the variability in site nitrogen availability (e.g. r 2 50.94 and 0.87 in 2001, when L was highest before declining due to droughts and storms), with the [CO2]-induced enhancement increasing with N (P 50.061). Incorporating data on N beyond the range of native fertility, achieved through N fertilization, indicated that pine L had reached the site maximum under elevated [CO2] where native N was highest. Thus closed canopy pine forests may be able to increase leaf area under elevated [CO2 ]i n moderate fertility sites, but are unable to respond to [CO2] in both infertile sites (insufficient resources) and sites having high levels of fertility (maximum utilization of resources). The total canopy L, representing the combined L of pine and hardwood species, was constant across the N gradient under both ambient and elevated [CO2], generating a constant enhancement of canopy L. Thus, in mixed species stands, L of canopy hardwoods which developed on lower fertility sites (� 3gNinputsm � 2 yr � 1 ) may be sufficiently enhanced under elevated [CO2] to compensate for the lack of response in pine L ,a nd generate an appreciable response of total canopy L (� 14%).

Abstract: We investigated microclimatic edge gradients associated with grassy powerlines, paved highways and perennial creeks in wet tropical forest in northeastern Australia during wet and dry seasons. Photosynthetically active radiation, air temperature and vapor pressure deficit, soil temperature, canopy temperature, soil moisture, and air speed in the rain forest understory were measured during traverses perpendicular to the forest edge. Light intensity was elevated near the edges of powerlines, highways, and creeks, but this effect was strongest for creek edges. Air temperature and vapor pressure deficit were elevated near powerline edges in the dry season and highway edges in both wet and dry seasons but were not elevated near creek edges in either season. In contrast, soil moisture was lowered near creek edges but not near either powerline or highway edges. No edge gradients were detected for air speed. Canopy temperature was elevated near highway edges and lowered near powerline edges in the wet season but no edge gradients in canopy temperature were detected near creek edges in either the wet or the dry season. We suggest that these different edge gradients may be largely the result of differences in the fluxes of latent and sensible heat within each type of linear canopy opening, with periodic flood disturbance assisting by maintaining a more open canopy near creek edges. Our data indicate that the nature of the linear canopy opening is at least as important as the width in determining the nature and severity of microclimatic edge effects, analogous to the "matrix effect" of traditional fragmentation studies.

Abstract: The application of detailed models of canopy photosynthesis rely on the estimation of attenuation of light in the canopy. This attenuation is readily estimated with the Lambert-Beer law when the canopy is homogeneous. In reality, forest canopies are far from homogeneous, and this has led to the use of detailed light extinction models that account for grouping of foliage between and within trees. Because such models require detailed parameterization and fine resolution inputs, they are impractical in larger-scale applications. Thus, there is interest in simplified models that can be readily parameterized. We developed two equations that can be used to estimate mean annual light interception by single unshaded trees and by stands of Poisson distributed trees. Interception by single trees is a function of crown surface area, the ratio of leaf area to crown surface area, the extinction coefficient in a homogeneous canopy--which can be determined separately--and one empirical parameter that depends on the mean solar angle. The summary model was tested against a detailed model of interception, and showed good agreement, although with slight bias. The results showed that crown surface area is a good summary variable for crown size and shape, because errors are independent of crown shape (ellipsoids, cones and height:width ratios). We also tested whether canopy photosynthesis is proportional to light interception across canopies differing in structure and leaf area index, and found that light-use efficiency is influenced by canopy structure. The model is useful in larger-scale applications because it can be parameterized with available data without the need for additional empirical parameters. It can also be used to study the effect of stand structure on mean annual light interception and productivity.

Abstract: This paper aims to assess the influence of canopy cover on lichen growth in boreal forests along a regional forest gradient. Biomass and area gain, and some acclimation traits, were assessed in the...

Abstract: In contrast to most insect guilds, gall-forming insects are thought to reach highest diversity on sclerophyllous vegetation, such as Neotropical savannas and Mediterranean vegetation types. The water and nutrient stress endured by meristems of canopy trees in tall wet tropical rainforests may cause leaf sclerophylly. Hence, the upper canopies of such ecosystems may represent a suitable habitat for gall-forming insects. At the San Lorenzo Protected Area, Panama, we estimated free-feeding herbivory and gall densities within five sites in 2003 and 2004, by surveying leaves in vertical and horizontal transects. In each sample, we recorded leaf density (mature and young foliage), free-feeding herbivore damage and number of galls, including the presence of live larvae, parasitoids or fungi. We surveyed 43 994 leaves, including 231 plants and 73 tree and liana species. We collected 5014 galls from 17 host-plant species, including 32 gall species of which 59% were restricted to the canopy (overall infestation rates: 2.4% in 2003, 5.5% in 2004). In 2003, 16% of the galls were occupied by live larvae, against 5% in 2004. About 1720% of leaves surveyed suffered from free-feeding herbivory. Leaf sclerophylly increased significantly with sampling height, while free-feeding herbivory decreased inversely. Conversely, the number of live galls collected in the canopy was 1316 times higher than in the understorey, a pattern consistent across sites and years. Hence, the probability of gall survivorship increased with increasing leaf sclerophylly as death by fungi, parasitoids or accidental chewing were greater in the understorey. Increasing harsh ecophysiological conditions towards the upper canopy appear favourable to galls-forming population maintenance, in support of the hypothesis of harsh environment. Hence, gall diversity and abundance in the upper canopy of tall tropical forests are perhaps among the highest in the world.