Yield development and nutrient dynamics in cocoa-gliricidia agroforests of Central Sulawesi, Indonesia

In the Napu and Palolo Valleys of Central Sulawesi, Indonesia, a chronosequence sought to identify the relationship between tree age, nutrient dynamics and cocoa (Theobroma cacao L.) yield in association with gliricidia (Gliricidia sepium (Jacq.) Steud.). The chronosequence surveyed cocoa-gliricidia plantations with a maximum age of 8 and 15 years, respectively, in Napu and Palolo. The characteristics of the valleys were also quite different, with an altitude of 1,139–1,166 m a.s.l. in Napu and 592–651 m a.s.l. in Palolo. Annual rainfall was 1,543 mm in Napu and 1,811 mm in Palolo. The yield of cocoa increased fairly steadily, with growth rates higher in Palolo than in Napu. Whereas a higher level of bean P led to a higher single bean weight (g d.w.) in Napu, a higher level of bean K led to a lower single bean weight in Palolo. The relatively high level of K appeared to have coincided with immature growth stages of cocoa. As trees matured, their increased rate of C assimilation was seen in the form of higher single bean weight. We found no statistically significant change in the soil’s carbon-nutrient levels when viewed over the entire timeframe of 8 and 15 years in the 2 valleys. In addition, there was no correlation between the soil’s carbon-nutrient levels and the single bean weight. Nor did we find any correlation between the soil’s carbon-nutrient levels impacting the bean’s carbon-nutrient levels. Of regression lines, P had the steepest slope and was considered the most limited nutrient relative to the other nutrients although its correlation was insignificant. The farmers’ estimation of cocoa yield was about 68% less than our measured bean weight per area per year (kg d.w. ha^?1 year^?1), implying a more refined definition of ripeness. In a cocoa agroforest, income could be supplemented by durable tree crops instead of growing gliricidia which is removed after several years of growth. This removal and the shallow rooting of cocoa indicate that the cocoa production would be sustainable only in the immediate future.     

Allometric relationships of frequently used shade tree species in cacao agroforestry systems in Sulawesi, Indonesia

Shade trees play an important role within agroforestry systems by influencing radiation and wind regimes as well as nutrient and hydrological cycling. However, there is a lack of quantitative assessments of their functions. One of the reasons is the rare information on structural characteristics of shade tree species. Therefore, the aim of this study is to provide basic information on the structure of frequently used shade tree species for the implementation of models simulating the ecosystem processes in agroforestry systems. The investigation of the shade trees was conducted at two cacao agroforestry sites on Sulawesi, Indonesia. The measurements of the main structural parameters: diameter at breast height, tree height, trunk height, crown length and crown radius were carried out for the shade tree species Aleurites moluccana, Cocos nucifera and Gliricidia sepium. For data collection, the National Forest Inventory Field Manual Template by FAO (2004) was applied. Based on this information allometric functions were derived for the correspondent shade tree species. The best significant relationships were obtained for the height-crown length relationship of the dicotyledonous tree species’ A. moluccana and G. sepium with a coefficient of determination r² = 0.925 and r² = 0.738, respectively, and the height-crown length relationship of the monocotyledonous palm C. nucifera with r² = 0.663. The transferability tests ‘analysis of covariance’ and ‘homogeneity of slopes’ have shown that the obtained allometric functions are also applicable to other cacao agroforestry systems of the region.     

The forest garden system of Saparua island Central Maluku, Indonesia, and its role in maintaining tree species diversity

The dusuns of Central Maluku which improve fallows by planting spice trees such as clove (Syzygium aromaticum syn. Eugenia aromatica), nutmeg (Myristica fragrans), or coconut (Cocos nucifera) and tending spontaneous regeneration, are an example of a species-rich forest garden system in the eastern hemisphere of the Indonesian archipelago. In this paper we report the influence of site conditions, as slope inclination, soil type, and soil depth, on species composition, tree species diversity, and stand structure in old dusuns in four villages on Saparua, a small island south of Seram. In addition, we compared the floristic composition and stand structure of old dusuns with those of primary forest and advanced fallow vegetation (∼15-yr old). Soil conditions, particularly the depth of the mineral soil layer, and the relief have a major influence on species selection in dusuns. The most common species in old dusuns were absent both in primary forest and forest fallow. Floristic similarity between old dusuns and the unmanaged stands was low. However, in two study villages species richness of old dusuns was similar to that of the primary forest. Old dusuns showed an open structure with an average reduction of basal area by about half compared to the primary forest. Less than half of all individuals found above 10 cm DBH in old dusuns were planted. Only very few species were exclusively planted. Most species were both planted and emerged from spontaneous regeneration. The results indicate the important role of spontaneous regeneration in this forest garden system. This revised version was published online in June 2006 with corrections to the Cover Date.     

Wood and non-wood forest products of Central Java,Indonesia

We investigated herb and woody species at rehabilitated forests planted by mahogany and teak, and original, not rehabilitated forests in Yogyakarta, Indonesia. Plant species were classified into five use categories: medicine, food, fodder, ornament, and construction. We registered 142 species belonging to 54 families known as useful species with at least one, often more, use categories. The number of useful species was highest for medicine use (107 species). There was a dominance of exotic herb species used for food and annual herbs used for fodder. The number of useful herbs and exotic woody species was highest in teak stands. The number of woody species used for medicine, food, and construction was higher than those for ornament and fodder. All herb species decreased with time, except annual native, that increased. Around 50% of the useful species occurred only once in one site, and some species showed a distribution restricted to one type of stands. Overall, our results seem to show that rehabilitated stands are doing well with regard to useful herb and woody species. We suggest strategies for plant and ecosystem conservation, especially for rare native plant species and species with restricted distribution.     

A link between hurricane-induced tree sprouting, high stem density and short canopy in tropical dry forest

The physiognomy of Caribbean dry forest is shorter, denser and contains a greater proportion of multi-stemmed trees than other neotropical dry forests. Our previous research, conducted after Hurricane Georges in 1998, has shown that dry forest trees sprout near the base following hurricane disturbance, even if the trees have not incurred structural damage. However, for these hurricane-induced sprouts to contribute to the physiognomy of the forest, they must grow and survive. We followed sprout dynamics and stem mortality on 1,407 stems from 1998, after Hurricane Georges, until 2005. The number of surviving sprouts and the proportion of sprouting stems decreased during the 7-year period, but the sprouting rate was still 3-fold higher and the proportion of sprouting stems 5-fold higher than before the hurricane. Mortality of non-sprouting stems (15.4%) was about the same as for sprouting stems (13.9%) after 7 years. The mean length of the dominant sprout surpassed 1.6 m by 2005, with over 13% of the dominant sprouts reaching subcanopy height. Sprout growth and survival varied among species. These results demonstrate that, despite some thinning, hurricane-induced sprouts survive and grow and that the unique physiognomic characteristic of Caribbean dry forests is related to hurricane disturbance.     

Photosynthesis of a tropical canopy tree, Ceiba pentandra, in a lowland forest in Panama

Diel (24 h) courses of CO(2) and water-vapor exchange of Ceiba pentandra (L.) Gaertn. (Bombacaceae) were studied under natural tropical conditions in the semi-evergreen moist forest of Barro Colorado Island, Panama. Measurements were conducted from early February 1991 (dry season), shortly after new leaves emerged, until mid-October 1991 (wet season), when leaves were shed. Rates of net CO(2) uptake were significantly higher in the dry season than in the wet season, and showed a linear decrease with leaf age. Leaf nitrogen concentrations and contents also decreased with age. Our estimate of annual carbon gain (2640 g CO(2) m(-2) year(-1) or 21 g CO(2) g(DW) (-1) year(-1)) is considerably higher than estimates available for temperate forest trees.     

Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain

? Interspecific differences in tree growth patterns with respect to biotic and abiotic factors are key for understanding forest structure and dynamics, and predicting potential changes under climate change.

? Repeated observations from the Spanish Forest Inventory (SFI) were used to parameterize maximum likelihood estimators of tree growth as a function of tree size, competition indices and climate for Pinus pinaster, P. sylvestris, Quercus ilex and Q. pyrenaica.

? Significant responses to both biotic and abiotic factors were found, with interspecific differences in species performance along competition, temperature and precipitation gradients. Q. ilex was the species most tolerant to competition while P. pinaster was the species most sensitive to climatic variation. Species relative positions shifted along gradients of these factors with rank reversals in species performance along size, competition and climatic gradients.

? The results based on average growth matched previous forestry classifications and experimental studies on relative growth rate (RGR).

? When examining growth along studied abiotic and biotic gradients, a mismatch was found between species performance ranks as predicted by our models and information derived from previous knowledge. Those discrepancies highlight the relevance of ontogeny and environmental heterogeneity in defining species performance along competition gradients.

     

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.     

Variation in photosynthetic light-use efficiency in a mountainous tropical rain forest in Indonesia

Photosynthetically active radiation (Q)-use efficiency (epsilon) is an important parameter for deriving carbon fluxes between forest canopies and the atmosphere from meteorological ground and remote sensing data. A common approach is to assume gross primary production (P(g)) and net primary production (P(n)) are proportional to Q absorbed by vegetation (Q(abs)) by defining the proportionality constants epsilon(Pg) and epsilon(Pn) (for P(g) and P(n), respectively). Although remote sensing and climate monitoring provide Q(abs) and other meteorological data at the global scale, information on epsilon is particularly scarce in remote tropical areas. We used a 16-month continuous CO(2) flux and meteorological dataset from a mountainous tropical rain forest in central Sulawesi, Indonesia to derive values of epsilon(Pg) and to investigate the relationship between P(g) and Q(abs). Absorption was estimated with a 1D SVAT model from measured canopy structure and short wave radiation. The half-hourly P(g) data showed a saturation response to Q(abs). The amount of Q(abs) required to saturate P(g) was reduced when water vapor saturation deficit (D) was high. Light saturation of P(g) was still evident when shifting from half-hourly to daily and monthly time scales. Thus, for a majority of observations, P(g) was insensitive to changes in Q(abs). A large proportion of the observed seasonal variability in P(g) could not be attributed to changes in Q(abs) or D. Values of epsilon(Pg) varied little around the long-term mean of 0.0179 mol CO(2) (mol photon)(-1) or 0.99 g C MJ(-1) (the standard deviations were +/- 0.006 and +/- 0.0018 mol CO(2) (mol photon)(-1) for daily and monthly means, respectively). In both cases, epsilon(Pg) values were more sensitive to Q(abs) than to daytime D. These findings show that the current epsilon-approaches fail to predict P(g) at our tropical rain forest site for two reasons: (1) they neglect saturation of P(g) when Q(abs) is high; and (2) they do not include factors, other than Q(abs) and D, that determine seasonality and annual sums of P(g).     

Impact of patch size on woody tree species richness and abundance in a tropical montane evergreen forest patches of south India

We examined the effects of forest patch size on woody tree species richness and abundance in tropical montane evergreen forest patches of the Nilgiri region,south India. We sampled woody trees(1 cm dbh) from 21 forest patches in the upper Nilgiri hills( 2000 m elevation) and recorded a total of 35,146 individuals of 61 species, 45 genera and 30 families. Species richness and abundance of sapling/shrubs(≥ 1 to 10 cm dbh)increased significantly with increasing patch size, but the species richness and abundance of small, medium and larger trees(≥ 10 to 30, ≥ 30 to 60 and ≥ 60 cm dbh,respectively) did not. Overall, forest interior species richness and abundance increased significantly with increasing patch size but edge species richness did not. Species richness and abundance of shade-tolerant and shade-demanding tree species also increased with increasing patch size. The abundance of zoochory dispersed tree species was significantly related to increasing patch size, but those dispersed by autochory did not display any clear relationship between patch size and species richness or abundance.Our findings suggest that with increasing forest patch area,tree compositional patterns may be driven by species specific shade-tolerance adaptations and dispersal patterns.Differential responses in these traits by the plant community within the individual habitat zones of forest edge and interiors likely plays a major role in determining the inherent plant community and thus the subsequent ecological processes of forest patches, including their responses to increasing patch area.     

Carbon stock measurements of a degraded tropical logged-over secondary forest in Manokwari Regency, West Papua, Indonesia

Several studies have been conducted in the past on carbon stock measurements in the tropical forests of Indonesia.This study is the first related research conducted in the New Guinea Island.In a degraded logged-over secondary forest in Manokwari Regency (West Papua,Indonesia),carbon stocks were measured for seven parts,i.e.,above-ground biomass (AGB),below-ground biomass (BGB),under-storey biomass (B u),necromass of dead leaves (N l),necromass of dead trees (N t),litter (L) and soil (S) using appropriate equations and laboratory analysis.Total carbon stocks were measured at 642.8 tC·ha-1 in the low disturbance area,536.9 tC·ha-1 in the moderate disturbance area and 490.4 tC·ha-1 in the high disturbance area.B u,N l and N t were not significant in the carbon stock and were collectively categorized as a total biomass complex.The carbon stock of litter was nearly equal to that of the total biomass complex,while the total carbon stock in the soil was eight times larger than the total biomass complex or the carbon stock of the litter.We confirmed that the average ratio of AGB and BGB to the total biomass (TB) was about 84.7% and 15.3%,respectively.Improvements were made to the equations in the low disturbance logged-over secondary forest area,applying corrections to the amounts of biomass of sample trees,based on representative commercial trees of category one.TB stocks before and after correction were estimated to be 84.4 and 106.7 tC·ha-1,indicating that these corrections added significant amounts of tree biomass (26.4%) during the sampling procedure.In conclusion,the equations for tree biomass developed in this study,will be useful for evaluating total carbon stocks,especially TB stocks in logged-over secondary forests throughout the Papua region.     

Radial variation in sap flow in five laurel forest tree species in Tenerife,Canary Islands

Variations in radial patterns of xylem water content and sap flow rate were measured in five laurel forest tree species (Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng., Myrica faya Ait., Erica arborea L. and Ilex perado Ait. ssp. platyphylla (Webb & Berth.) Tutin) growing in an experimental plot at Agua García, Tenerife, Canary Islands. Measurements were performed around midday during warm and sunny days by the heat field deformation method. In all species, water content was almost constant (around 35% by volume) over the whole xylem cross-sectional area. There were no differences in wood color over the whole cross-sectional area of the stem in most species with the exception of E. arborea, whose wood became darker in the inner layers. Radial patterns of sap flow were highly variable and did not show clear relationships with tree diameter or species. Sap flow occurred over the whole xylem cross-sectional area in some species, whereas it was limited to the outer xylem layers in others. Sap flow rate was either similar along the xylem radius or exhibited a peak in the outer part of the xylem area. Low sap flow rates with little variation in radial pattern were typical for shaded suppressed trees, whereas dominant trees exhibited high sap flow rates with a peak in the radial pattern. Stem damage resulted in a significant decrease in sap flow rate in the outer xylem layers. The outer xylem is more important for whole tree water supply than the inner xylem because of its larger size. We conclude that measurement of radial flow pattern provides a reliable method of integrating sap flow from individual measuring points to the whole tree.     

Radial profiles of sap flow with increasing tree size in maritime pine

We investigated the radial variation of sap flow within sapwood below the live crown in relation to tree size in 10-, 32-, 54- and 91-year-old maritime pine stands (Pinus pinaster Ait.). Radial variations were determined with two thermal dissipation sensors; one measured sap flux in the outer 20 mm of the xylem (Jref), whereas the other was moved radially across the sapwood in 20-mm increments to measure sap flux at multiple depths (Jref). For all tree sizes, sap flow ratios (Ri = JiJref (-1)) declined with increasing sapwood depth, but the decrease was steeper in trees with large diameters. Correction factors (C) were calculated to extrapolate Jref for an estimate of whole-tree sap flux. A negative linear relationship was established between stem diameter and C, the latter ranging from 0.6 to 1.0. We found that neglecting these radial corrections in 10-, 32-, 54- and 91-year-old trees would lead to overestimation of stand transpiration by 4, 14, 26 and 47%, respectively. Therefore, it is necessary to account for the differential radial profiles of sap flow in relation to tree size when comparing tree transpiration and hydraulic properties among trees differing in size.     

Below- and above-ground biomass and net primary production in a paleotropical natural forest (Sulawesi,Indonesia) as compared to neotropical forests

Data on the biomass and productivity of southeast Asian tropical forests are rare, making it difficult to evaluate the role of these forest ecosystems in the global carbon cycle and the effects of increasing deforestation rates in this region. In particular, more precise information on size and dynamics of the root system is needed. In six natural forest stands at pre-montane elevation (c. 1000 m a.s.l.) on Sulawesi (Indonesia), we determined above-ground biomass and the distribution of fine (d < 2 mm) and coarse roots (d > 2 mm), estimated above- and below-ground net production, and compared the results to literature data from other pre-montane paleo- and neotropical forests. The mean total biomass of the stands was 303 Mg ha⁻¹ (or 128 Mg C ha⁻¹), with the largest biomass fraction being recorded for the above-ground components (286 Mg ha⁻¹) and 11.2 and 5.6 Mg ha⁻¹ of coarse and fine root biomass (down to 300 cm in the soil profile), resulting in a remarkably high shoot:root ratio of c. 17. Fine root density in the soil profile showed an exponential decrease with soil depth that was closely related to the concentrations of base cations, soil pH and in particular of total P and N. The above-ground biomass of these stands was found to be much higher than that of pre-montane forests in the Neotropics, on average, but lower compared to other pre-montane forests in the Paleotropics, in particular when compared with dipterocarp forests in Malesia. The total above- and below-ground net primary production was estimated at 15.2 Mg ha⁻¹ yr⁻¹ (or 6.7 Mg C ha⁻¹ yr⁻¹) with 14% of this stand total being invested below-ground and 86% representing above-ground net primary production. Leaf production was found to exceed net primary production of stem wood. The estimated above-ground production was high in relation to the mean calculated for pre-montane forests on a global scale, but it was markedly lower compared to data on dipterocarp forests in South-east Asia. We conclude that the studied forest plots on Sulawesi follow the general trend of higher biomasses and productivity found for paleotropical pre-montane forest compared to neotropical ones. However, biomass stocks and productivity appear to be lower in these Fagaceae-rich forests on Sulawesi than in dipterocarp forests of Malesia.     

Timber tree regeneration along abandoned logging roads in a tropical Bolivian forest

Sustainable management of selectively logged tropical forests requires that felled trees are replaced through increased recruitment and growth. This study compares road track and roadside regeneration with regeneration in unlogged and selectively logged humid tropical forest in north-eastern Bolivia. Some species benefited from increased light intensities on abandoned logging roads. Others benefited from low densities of competing vegetation on roads with compacted soils. This was the case for the small-seeded species Ficus boliviana C.C. Berg and Terminalia oblonga (Ruiz & Pav.) Steud. Some species, e.g. Hura crepitans L., displayed patchy regeneration coinciding with the presence of adult trees. Our results suggest that current management practices could be improved by intensifying logging in some areas to improve regeneration of light demanding species. Sufficient seed input in logged areas should be ensured by interspersing large patches of unlogged forest with logged areas. This may also assist regeneration of species that perform poorly in disturbed areas.     

Stomatal conductance, transpiration and sap flow of tropical montane rain forest trees in the southern Ecuadorian Andes

We investigated tree water relations in a lower tropical montane rain forest at 1950-1975 m a.s.l. in southern Ecuador. During two field campaigns, sap flow measurements (Granier-type) were carried out on 16 trees (14 species) differing in size and position within the forest stand. Stomatal conductance (g(s)) and leaf transpiration (E(l)) were measured on five canopy trees and 10 understory plants. Atmospheric coupling of stomatal transpiration was good (decoupling coefficient Omega = 0.25-0.43), but the response of g(s) and E(l) to the atmospheric environment appeared to be weak as a result of the offsetting effects of vapor pressure deficit (VPD) and photosynthetic photon flux (PPF) on g(s). In contrast, sap flow (F) followed these atmospheric parameters more precisely. Daily F depended chiefly on PPF sums, whereas on short time scales, VPD impeded transpiration when it exceeded a value of 1-1.2 kPa. This indicates an upper limit to transpiration in the investigated trees, even when soil water supply was not limiting. Mean g(s) was 165 mmol m(-2) s(-1) for the canopy trees and about 90 mmol m(-2) s(-1) for the understory species, but leaf-to-leaf as well as tree-to-tree variation was large. Considering whole-plant water use, variation in the daily course of F was more pronounced among trees differing in size and crown status than among species. Daily F increased sharply with stem diameter and tree height, and ranged between 80 and 120 kg day(-1) for dominant canopy trees, but was typically well below 10 kg day(-1) for intermediate and suppressed trees of the forest interior.     

Biomass of a man-made forest of timber tree species in the humid tropics of West Java, Indonesia

Biomass of a mature man-made forest in West Java, Indonesia, was estimated to evaluate the carbon sequestration potential of plantation forest in the humid tropics. Twenty plots, each 0.25 ha in area and containing one to six planted species over 40 years of age and with closed canopies, were selected. Trunk dry mass was estimated from trunk diameter, tree height, and bulk density. Maximum trunk diameter (122 cm) was observed in a 46-year-old Khaya grandifoliola C. DC. tree, and the tallest tree (51 m) was a 46-year-old Shorea selanica (DC.) Blume. The largest trunk biomass (911 Mg ha⁻¹) was achieved in the plot composed of two Khaya spp. Among the plots composed of indigeneous Dipterocarpaceae species, the largest trunk biomass was 635 Mg ha⁻¹. These trunk biomasses were larger than those reported from primary rainforests in Southeast Asia (e.g., 403 Mg ha⁻¹ in East Kalimantan, 522 and 368 Mg ha⁻¹ in Peninsular Malaysia). The large biomass in this forest suggests that, given favorable conditions, man-made forests can accumulate the quantities of atmospheric carbon that were lost by the logging of primary forests in the humid tropics.     

Stand-scale transpiration estimates in a Moso bamboo forest: (I) Applicability of sap flux measurements

The applicability of sap flux (F_d) measurements to bamboo forests has not been studied. This study was undertaken to establish an optimal and effective design for stand-scale transpiration (E) estimates in a Moso bamboo forest. To this aim, we validated F_d measurements in Moso bamboos in a cut bamboo experiment. In addition, we analyzed how sample sizes affect the reliability of E estimates calculated from F_d and conducting culm area (A_{S_b}). In the cut bamboo experiments, we found that F_d measurement using a 10 mm probe was a valid means of determining the water-use behavior of a Moso bamboo, although a specific correction was needed. Furthermore, we calculated E from stand A_{S_b} (A_{S_stand}) and mean stand F_d (J_S). Employing Monte Carlo analysis, we examined potential errors associated with sample size in E, A_{S_stand}, and J_S using an original dataset with A_{S_b} and F_d measured for 40 and 16 individuals, respectively. Consequently, we determined the optimal sample size for both A_{S_stand} and J_S estimates as 11. The optimal sample sizes for J_S were almost the same under different vapor pressure deficit and soil moisture conditions. The optimal sample size for J_S at the study site was less than that of a coniferous plantation in the same region probably owing to small individual-to-individual variations in sap flux in the Moso bamboo forest. Our study concludes that sap flux measurements are an applicable technique for assessing water use in Moso bamboo forests.