Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d -1 in a 0·20-mdiameter individual of Cecropia longipes to 54 kg d -1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9-15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m 2 increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.
Extensive areas of the tropics have been converted into pasture for cattle ranching. Frequently, abandoned pasture does not revert to forest. The goal of this project was to identify barriers to lowland moist forest regeneration in highly degraded grasslands in the Sierra Nevada de Santa Marta, Colombia. The barriers we considered were seed source, seed predation, competition with grasses, microclimate and soil limitations on plant growth, and fire. Seed dispersal into the grasslands is limited to within 10 meters of forest fragments, but this barrier can be overcome by sowing seeds and planting seedlings and by establishing perches to attract dispersers. In these degraded grasslands, seed predation was lower than in the adjacent forest patches, and there was no evidence that grasses inhibited the establishment of woody species. The most important barrier was the severe degradation of the soils. In much of the area, the A and B horizons have been eroded away, leaving saprolite at the soil surface. Seedlings of two fast‐growing pioneer species, Ochroma pyramidale and Cochlospermum vitifolium, grew to a maximum height of only 2.5 and 12 cm, respectively, during the first eight months. The slow plant growth in the degraded grassland soils compared to forest soils was associated with lower levels of cation‐exchange capacity, calcium, magnesium, and potassium. Even if these barriers could be overcome, the frequent and extensive use of fire in the region must be controlled to avoid killing established woody plants.
Little is known about partitioning of soil water resources in species-rich, seasonally dry tropical forests. We assessed spatial and temporal patterns of soil water utilization in several canopy tree species on Barro Colorado Island, Panama, during the 1997 dry season. Stable hydrogen isotope composition (δD) of xylem and soil water, soil volumetric water content (θ), and sap flow were measured concurrently. Evaporative fractionation near the soil surface caused soil water δD to decrease from about -15‰ at 0.1 m to -50 to -55‰ at 1.2 m depth. Groundwater sampled at the sources of nearby springs during this period yielded an average δD value of -60‰. θ increased sharply and nearly linearly with depth to 0.7 m, then increased more slowly between 0.7 and 1.05 m. Based on xylem δD values, water uptake in some individual plants appeared to be restricted largely to the upper 20 cm of the soil profile where θ dropped below 20% during the dry season. In contrast, other individuals appeared to have access to water at depths greater than 1 m where θ remained above 45% throughout the dry season. The depths of water sources for trees with intermediate xylem δD values were less certain because variation in soil water δD between 20 and 70 cm was relatively small. Xylem water δD was also strongly dependent on tree size (diameter at breast height), with smaller trees appearing to preferentially tap deeper sources of soil water than larger trees. This relationship appeared to be species independent. Trees able to exploit progressively deeper sources of soil water during the dry season, as indicated by increasingly negative xylem δD values, were also able to maintain constant or even increase rates of water use. Seasonal courses of water use and soil water partitioning were associated with leaf phenology. Species with the smallest seasonal variability in leaf fall were also able to tap increasingly deep sources of soil water as the dry season progressed. Comparison of xylem, soil, and groundwater δD values thus pointed to spatial and temporal partitioning of water resources among several tropical forest canopy tree species during the dry season.
Distribution patterns of vascular plants with diameter at breast height (dbh) ≥ 2.5 cm were studied on the basis of compositional data from 30 small plots located in a rain-forest area in Colombian Amazonia. The research questions were: How are distribution patterns of species in relation to local abundance in plots? Do understorey species (defined as species with individuals that never attained dbh ≥ 10 cm anywhere) show better correlations with soils and environment than canopy species (defined as species with individuals that attained dbh ≥ 10 cm)? Are patterns found in the entire range of landscape units comparable to those found in well-drained uplands alone? Species that occurred in more than one plot showed higher local abundances. This pattern was consistent among environmental generalists and specialists. Locally rare species (with only one individual in a plot) occurred mostly in well-drained uplands. Considering all landscape units, Mantel tests showed substantial correlations between environmental data (soil chemical data, drainage and flooding) and species composition. Canopy species were only slightly less correlated with environmental data than understorey species. Elimination of the spatial component in the data did not reduce these correlations. In well-drained uplands, understorey species were better correlated with soils than canopy species. Here, however, the spatial configuration of the plots became more important in explaining species patterns.
Source water used by plants of several species in a semi-evergreen lowland tropical forest on Barro Colorado Island, Panama, was assessed by comparing the relative abundance of deuterium, D, versus hydrogen, H (stable hydrogen isotope composition, δD) in xylem sap and in soil water at different depths, during the dry season of 1992. Ecological correlates of source water were examined by comparing xylem water δD values with leaf phenology, leaf water status determined with a pressure chamber, and rates of water use determined as mass flow of sap using the stem heat balance method. Soil water δD values decreased sharply to 30 cm, then remained relatively constant with increasing depth. Average δD values were-13‰, for 0-30 cm depth and-36.7‰ for 30-100 cm depth. Soil water δD values were negatively associated with soil water content and soil water potential. Concurrent analyses of xylem water revealed a high degree of partitioning of water resources among species of this tropical forest. Xylem water δD of deciduous trees (average=-25.3±1.4‰) was higher than that of evergreen trees (average=-36.3±3.5‰), indicating that evergreen species had access to the more abundant soil water at greater depth than deciduous species. In evergreen shade-tolerant and high-light requiring shrubs and small trees, δD of xylem water was negatively correlated with transpiration rate and leaf water potential indicating that species using deeper, more abundant water resources had both higher rates of water use and more favorable leaf water status.
Environmental and physiological regulation of transpiration were examined in several gap-colonizing shrub and tree species during two consecutive dry seasons in a moist, lowland tropical forest on Barro Colorado Island, Panama. Whole plant transpiration, stomatal and total vapor phase (stomatal + boundary layer) conductance, plant water potential and environmental variables were measured concurrently. This allowed control of transpiration (E) to be partitioned quantitatively between stomatal (g ) and boundary layer (g) conductance and permitted the impact of invividual environmental and physiological variables on stomatal behavior and E to be assessed. Wind speed in treefall gap sites was often below the 0.25 m s stalling speed of the anemometer used and was rarely above 0.5 m s, resulting in uniformly low g (c. 200-300 mmol m s) among all species studied regardless of leaf size. Stomatal conductance was typically equal to or somewhat greater than g . This strongly decoupled E from control by stomata, so that in Miconia argentea a 10% change in g when g was near its mean value was predicted to yield only a 2.5% change in E. Porometric estimates of E, obtained as the product of g and the leaf-bulk air vapor pressure difference (VPD) without taking g into account, were up to 300% higher than actual E determined from sap flow measurements. Porometry was thus inadequate as a means of assessing the physiological consequences of stomatal behavior in different gap colonizing species. Stomatal responses to humidity strongly limited the increase in E with increasing evaporative demand. Stomata of all species studied appeared to respond to increasing evaporative demand in the same manner when the leaf surface was selected as the reference point for determination of external vapor pressure and when simultaneous variation of light and leaf-air VPD was taken into account. This result suggests that contrasting stomatal responses to similar leaf-bulk air VPD may be governed as much by the external boundary layer as by intrinsic physiological differences among species. Both E and g initially increased sharply with increasing leaf area-specific total hydraulic conductance of the soil/root/leaf pathway (G ), becoming asymptotic at higher values of G. For both E and g a unique relationship appeared to describe the response of all species to variations in G. The relatively weak correlation observed between g and midday leaf water potential suggested that stomatal adjustment to variations in water availability coordinated E with water transport efficiency rather than bulk leaf water status.
Concurrent, independent measurements of stomatal conductance (g^), transpiration [E) and microenvironmental variables were used to characterize control of crown transpiration in four tree species growing in a moist, lowland tropical forest. Access to the upper forest canopy was provided by a construction crane equipped with a gondola. Estimates of boundary layer conductance (g,,) obtained with two independent methods permitted control of E to be partitioned quantitatively between gâ nd gh using ^ dimensionless decoupling coefficient (12) ranging from zero to 1. A combination of high g^ (c. 300-600 mmol m~^ s~') and low wind speed, and therefore relatively low g,, (c. 100-800 mmol m~^ s"'), strongly decoupled E from control by stomata in all four species (Q = 0-7-0-9). Photosyntbetic water-use efficiency was predicted to increase rather than decrease with increasing gs because g,, was relatively low and internal conductance to CO2 transfer was relatively high. Responses of g, to humidity were apparent only when the leaf surface, and not the bulk air, was used as the reference point for determination of external vapour pressure. However, independent measurements of crown conductance (g^.), a total vapour phase conductance that included stomatal and boundary layer components, revealed a clear decline in ge with increasing leaf-to-bulk air vapour pressure difference (V^), because the external reference points for determination of ge and V,, were compatible. The relationships between g^ and V,, and between g, and V, appeared to be distinct for each species. However, when gj, and gj were normalized by the branch-specific ratio of leaf area to sapwood area (LA/SA), a morphological index of potential transpirational demand relative to water transport capacity, a common relationship between conductance and evaporative demand for all four species emerged. Taken together, these results implied that, at a given combination of LA/SA and evaporative demand scaled to the appropriate reference point, the vapour phase conductance and therefore transpiration rates on a leaf area basis were identical in all four contrasting species studied.
Stomatal control of crown transpiration was studied in Anacardium excelsum, a large‐leaved, emergent canopy species common in the moist forests of Central and northern South America. A construction crane equipped with a gondola was used to gain access to the uppermost level in the crown of a 35‐m‐tall individual. Stomatal conductance at the single leaf scale, and transpiration and total vapour phase conductance (stomatal and boundary layer) at the branch scale were measured simultaneously using the independent techniques of porometry and stem heat balance, respectively. This permitted the sensitivity of transpiration to a marginal change in stomatal conductance to be evaluated using a dimensionless coupling coefficient (1‐ω) ranging from zero to 1, with 1 representing maximal stomatal control of transpiration. Average stomatal conductance varied from 0.09 mol m−2 s−1 during the dry season to 0.3 mol m−2 s−1 during the wet season. Since boundary layer conductance was relatively low (0.4 mol m−2 s−1), 1‐ω ranged from 0.46 during the dry season to only 0.25 during the wet season. A pronounced stomatal response to humidity was observed, which strongly limited transpiration as evaporative demand increased. The stomatal response to humidity was apparent only when the leaf surface was used as the reference point for measurement of external vapour pressure. Average transpiration was predicted to be nearly the same during the dry and wet seasons despite a 1 kPa difference in the prevailing leaf‐to‐air vapour pressure difference. The patterns of stomatal behaviour and transpiration observed were consistent with recent proposals that stomatal responses to humidity are based on sensing the transpiration rate itself.
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