Litter Decomposition and Nutrient Release in a Tropical Seasonal Rain Forest of Xishuangbanna,Southwest China1

We studied litter decomposition and nutrient release in a tropical seasonal rain forest of Xishuangbanna, Southwest China. The monthly decay rates (k) of leaf litter ranged from 0.02 to 0.21/mo, and correlated with rainfall and soil moisture. Annual k values for leaf litter (1.79/yr) averaged 4.2 times of those for coarse wood (2.5–3.5 cm in diameter). The turnover coefficients of forest floor mass (annual litterfall input/mean floor mass) were: 4.11/yr for flowers and fruits, 2.07/yr for leaves, and 1.17/yr for fine wood (≤2 cm in diameter), with resident time decreasing from fine woods (0.85 yr) to leaves (0.48 yr) and to flower and fruits (0.24 yr). Nutrient residence times in the forest floor mass were ranked as: Ca (1.0 yr) > P (0.92 yr) > Mg (0.64 yr) > N (0.36 yr) > K (0.31 yr). Our data suggest that rates of litter decomposition and nutrient release in the seasonal rain forest of Xishuangbanna are slower than those in typical lowland rain forests, but similar to those in tropical semideciduous forests.     

Spatial Variability in Litterfall,Litter Standing Crop and Litter Quality in a Tropical Rain Forest Region

Understanding the spatial variability in plant litter processes is essential for accurate comprehension of biogeochemical cycles and ecosystem function. We assessed spatial patterns in litter processes from local to regional scales, at sites throughout the wet tropical rain forests of northern Australia. We aimed to determine the controls (e.g., climate, soil, plant community composition) on annual litter standing crop, annual litterfall rate and in situ leaf litter decomposability. The level of spatial variance in these components, and leaf litter N, P, Ca, lignin, α‐cellulose and total phenolics, was determined from within the scale of subregion, to site (1 km transects) to local/plot (~30 m²). Overall, standing crop was modeled with litterfall and its chemical composition, in situ decomposability, soil Na, and topography (r² = 0.69, 36 plots). Litterfall was most closely aligned with plant species richness and stem density (negative correlation); leaf decomposability with leaf‐P and lignin, soil Na, and dry season moisture (r² = 0.89, 40 plots). The predominant scale of variability in litterfall rates was local (plot), while litter standing crop and α‐cellulose variability was more evenly distributed across spatial scales. Litter decomposability, N, P and phenolics were more aligned with subregional differences. Leaf litter C, lignin and Ca varied most at the site level, suggesting more local controls. We show that variability in litter quality and decomposability are more easily accounted for spatially than litterfall rates, which vary widely over short distances possibly in response to idiosyncratic patterns of disturbance.     

Phenology and Stem Diameter Increment Seasonality in a Costa Rican Wet Tropical Forest

The relationship between phenology and tree stem diameter increment is largely unexplored in tropical species, especially in wet tropical forests. To explore links between these phenomena, we measured stem diameter increment and phenology of ten canopy tree species from a range of functional types in the Atlantic lowlands of Costa Rica to test for seasonal and interannual patterns. We measured stem diameter increment using band dendrometers and visually assessed leaf and reproductive phenology monthly from 1997 to 2000. We categorized the species into groups based on patterns of leaf exchange and reproduction. Species were either deciduous with synchronous or asynchronous leaf drop, or evergreen with continuous or seasonal leaf flushing. Flowering occurred supra-annually, annually, or continuously. Of the ten species studied, four species, Cecropia insignis, Dipteryx panamensis, Lecythis ampla, and Simarouba amara , had consistent seasonal stem diameter increment patterns in both years. Dipteryx panamensis and L. ampla were deciduous with synchronized leaf drop . Cecropia insignis was evergreen and produced new leaves continuously. Simarouba amara , also evergreen, exchanged leaves over a brief period once a year. We tested whether stem diameter increment was correlated to phenology using logistic regression. Leaflessness significantly explained patterns in stem diameter increment but reproductive phenology did not. Deciduous trees were 2.6–9.3 times more likely to grow less than average the month following leaffall than in months when trees had full crowns.     

Density, Distribution, and Attributes of Tree Cavities in an Old-Growth Tropical Rain Forest

Tree cavities are a critical resource for many animals, especially as nesting sites for birds. Patterns of cavity distribution in temperate forests are well studied, yet little is known of cavities in tropical forests, despite a hypothesized decrease in cavity availability with decreasing latitude. We studied cavity density and distribution in a wet lowland tropical forest in Costa Rica and compared our results with estimates from forests around the world. Cavities at our site were common, occurred frequently in living trees, and were often formed by damage or decay rather than by woodpeckers. Most cavities had small openings, and woodpecker-created cavities were nonrandomly oriented. Contrary to prediction, cavity density appears to increase from the poles to the tropics. We suggest potential mechanisms to explain these patterns.     

Tropical Rain Forest Structure,Tree Growth and Dynamics along a 2700-m Elevational Transect in Costa Rica

Rapid biological changes are expected to occur on tropical elevational gradients as species migrate upslope or go extinct in the face of global warming. We established a series of 9 1-ha plots in old-growth tropical rainforest in Costa Rica along a 2700 m relief elevational gradient to carry out long-term monitoring of tropical rain forest structure, dynamics and tree growth. Within each plot we mapped, identified, and annually measured diameter for all woody individuals with stem diameters >10 cm for periods of 3-10 years. Wood species diversity peaked at 400-600 m and decreased substantially at higher elevations. Basal area and stem number varied by less than two-fold, with the exception of the 2800 m cloud forest summit, where basal area and stem number were approximately double that of lower sites. Canopy gaps extending to the forest floor accounted for <3% of microsites at all elevations. Height of highest crowns and the coefficient of variation of crown height both decreased with increasing elevation. Rates of turnover of individuals and of stand basal area decreased with elevation, but rates of diameter growth and stand basal area showed no simple relation to elevation. We discuss issues encountered in the design and implementation of this network of plots, including biased sampling, missing key meteorological and biomass data, and strategies for improving species-level research. Taking full advantage of the major research potential of tropical forest elevational transects will require sustaining and extending ground based studies, incorporation of new remotely-sensed data and data-acquisition platforms, and new funding models to support decadal research on these rapidly-changing systems.     

Asymbiotic Nitrogen Fixation and Litter Decomposition on a Long Soil-Age Gradient in Hawaiian Montane Rain Forest1

We determined rates of decomposition and asymbiotic nitrogen fixation in the leaf litter of Cheirodendron spp. on the Hawaiian Islands. Leaf litter was collected from four sites on a long soil-age gradient (300 yr to 4.1 M yr) and decomposed at two sites that differed widely in substrate age and nutrient availability. Rates of decomposition were higher in litter decomposed at the older site, where nutrient availability was greater. A substantial amount of nitrogen and phosphorus immobilization occurred in litter decomposed at the older site, with more immobilization occurring in litter with lower initial nitrogen and phosphorus concentrations, suggesting both supply and demand controls on nutrient immobilization. Potential rates of nitrogen fixation were very low in the first 25 d (0–5 nmol acetylene/gdw/h), rose to much higher rates by 70 d (20–45 nmol), and then declined by 140 d. We found no significant difference in rates of potential nitrogen fixation between sites of decomposition, but there was a strong substrate effect, with higher rates in litter with low lignin, low nitrogen, and high phosphorus. Where significant immobilization of nitrogen occurred for decomposing Cheirodendron, nitrogen fixation could have comprised no more than 10 percent of immobilized nitrogen. Overall, rates of nitrogen fixation were dependent on the source of the decomposing substrate but not on the site of decomposition, while short-term decomposition and nutrient immobilization were strongly dependent on the site of decomposition but not as much on the source of the decomposing substrate.     

Winds under the Rain Forest Canopy: The Aerodynamic Environment of Gliding Tree Frogs

Movement of air under the canopy of a forest affects the gliding of animals such as frogs, snakes, geckos, and squirrels; the dispersal of pollen, seeds, and spores; as well as convective transport of heat and carbon dioxide. Wind speed profiles were measured under the canopy of a lowland rain forest during the morning, afternoon, and night at three sites in Costa Rica to determine the aerodynamic environment in which tree frogs maneuver while gliding. During the course of a day, average and maximum wind speeds were highest in the morning and midday, and lowest at night. Wind speeds under the canopy were highest near the top of the canopy and were lowest near the canopy floor in the morning and afternoon, and exhibited little variation with respect to height at night. Turbulence intensity (a common measure of gustiness) was constant (ca 1) for all times of day, heights in the canopy, and sites, but the absolute magnitudes of wind gust speeds were higher during the day than at night. Power spectral densities revealed that most of the variation in wind speeds occurred at frequencies that could potentially affect the gliding of tree frogs. Tree frogs (and many other gliding animals), however, glide at night and thereby avoid the higher wind speeds that occur by day. Computer simulations of the dynamic motions of frogs while gliding revealed that the night levels of wind gusts have little effect on the direction of gliding of tree frogs.     

Arthropod Abundance,Canopy Structure,and Microclimate in a Bornean Lowland Tropical Rain Forest1

This study applies a novel, vertically stratified fogging protocol to document arthropod abundance, density, and biomass across a vertical gradient in a primary, lowland dipterocarp forest canopy in Borneo. We fogged arthropods at 5 m vertical intervals and 20 m horizontal intervals along six full‐canopy transects and measured leaf surface areas along the same transects. The results show that arthropod biomass in the aboveground regions was 23.6 kg/ha, the abundance was 23.9 million individuals/ha, and the density on leaf surfaces was 280 individuals/m² leaf area. All three numbers are five to ten times higher than estimated by previous surveys of tropical lowland rain forest canopies using mass‐collection techniques. Arthropod abundance and biomass were analyzed in relation to canopy structure, composition, vapor pressure deficit (VPD), photosynthetic photon flux density (PPFD), and height. Using stepwise regression we found that 13 of 14 arthropod groups had significant positive relationships with one‐sided leaf area, 11 had significant negative relationships with VPD, 3 had significant relationships with height, and none showed positive relationships with light. Classifying the 14 taxa based on their responses to leaf area and VPD created three groups that corresponded roughly to the biology of these taxa. This study suggests that the biomass and abundance, and perhaps therefore—by extrapolation—the biodiversity, of tropical canopy arthropods may be very much higher than previously estimated.     

Seedling Establishment of a Canopy Tree Species in Malaysian Tropical Rain Forests

Abstract Unlike many other canopy tree species in tropical rain forests, Dryobalanops aromatica (Dipterocar-paceae, local name: Kapur) establishes monospecific dominant (monodominant) forests in Peninsular Malaysia. In natural conditions, monodominance of Kapur does not necessarily mean low species diversity of the Kapur forests. While the emergent canopy layer is occupied by Kapur, many other plant species, which are common to lowland dipterocarp primary forests in the same region, are found in lower canopy layer and understory.
To understand the ecological implications of the monodominance of Kapur, we monitored post dispersal survival and seedling establishment in a pure stand of Kapur in a plantation in Kepong, near Kuala Lumpur. Immediately after seed fall, seeds and cotyledon-stage seedlings suffered high predation by vertebrates such as rodents. The predation pressure was higher in a more general fruiting year (1991) than in a sporadic fruiting year (1992). In contrast to the high mortality of seeds and newly emerged seedlings, seedlings surviving to the six-leaf stage showed low mortality, which allowed the establishment of a sapling bank.
The occurrence of saplings of Kapur with a wide range of size classes in natural forests indicates that this species is more shade tolerant than other dipterocarp species such as the Shorea group and that it could well respond to enhanced light conditions caused by canopy opening. These characteristics may partly contribute to maintaining monodominance of Kapur.     

Leaf Litter Decomposition and Monodominance in the Peltogyne Forest of Marací Island, Brazil1

The forest type dominated by Peltogyne gracilipes (Caesalpiniaceae) on the riverine Marací Island is the least speciesrich of any recorded for Brazilian Amazonia. Because the forest has high soil and foliar Mg concentrations, and Mg is known to be toxic to plant growth at high concentrations, this study tested the hypothesis that dominance by Peltogyne is related to Mg leaf litter amounts and decomposition. We predicted that decomposition of Peltogyne leaves would differ from that of other species, and that their decomposition would result in a pulse of Mg release. Three plots (50 × 50 m) were established in each of three forest types: Peltogyne‐rich forest (PRF; dominated by P. gracilipes),Peltogyne‐poor forest (PPF), and forest without Peltogyne (FWP). Three leaf litter decomposition experiments tested if decomposition of mixed leaf litter in coarse‐ mesh (CM) litterbags differed among forests (experiment 1); whether or not decomposition and nutrient release of Ecclinusa guianensis, Lueheopsis duckeana, and Peltogyne in CM litterbags differed among forests and species (experiment 2); and using fine‐mesh (FM) litterbags, investigated the differences in the influence of fauna! activity on Ecclinusa and Peltogyne decomposition (experiment 3). Decomposition was independent of the presence and dominance of Peltogyne, since decomposition rates in both PRF and FWP were in general lower than in PPF. These differences appeared to be related to fauna] activity. The decomposition of Peltogyne leaves was lower than that of the other species tested and was more affected by microbial and physical action. It is possible that the monodominance of Peltogyne is related to its deciduousness and faster decomposition in the dry season, which coincides with a large leaf fall. Magnesium was lost quickly from the Peltogyne leaves and the resultant pulses of Mg into the soil during the heavy rains at the beginning of the wet season may be deleterious for other species that are not adapted to high solution Mg concentrations. Results obtained were consistent with the hypothesis that Peltogyne dominance is related to the pattern of its leaf decomposition and the seasonal pulses of toxic Mg.     

Foliar and Litter Nutrients, Nutrient Resorption, and Decomposition in Hawaiian Me t rosideros polymorpha

The native tree Metrosideros polymorpha dominates Hawaiian forests across a very wide range of soil fertility, including both sites where forest production is limited by nitrogen (N) and others where it is limited by phosphorus (P). Five long-term fertilization experiments have further broadened the range of nutrient availabilities experienced by Metrosideros. Adding P to P-limited sites increased foliar P concentrations threefold and litter P concentrations up to 10-fold; lignin concentrations decreased, and the decomposability of leaf litter increased from 32%–35% to 36%–46% mass loss in the first year. Adding N to N-limited sites increased leaf and litter N concentrations by only 15%–20%, with little or no effect on the decomposability of tissue. Received 22 January 1998; accepted 4 May 1998.     

Arthropod Abundance and Diversity in a Lowland Tropical Forest Floor in Panama: The Role of Habitat Space vs. Nutrient Concentrations

Tropical forest floor characteristics such as depth and nutrient concentrations are highly heterogeneous even over small spatial scales and it is unclear how these differences contribute to patchiness in forest floor arthropod abundance and diversity. In a lowland tropical forest in Panama we experimentally increased litter standing crop by removing litter from five plots (L−) and adding it to five other plots (L+); we had five control plots. After 32 mo of treatments we investigated how arthropod abundance and diversity were related to differences in forest floor physical (mass, depth, water content) and chemical properties (pH, nutrient concentrations). Forest floor mass and total arthropod abundance were greater in L+ plots compared with controls. There were no treatment differences in nutrient concentrations, pH or water content of the organic horizons. Over all plots, the mass of the fermentation horizon (Oe) was greater than the litter horizon (Oi); arthropod diversity and biomass were also greater in the Oe horizon but nutrient concentrations tended to be higher in the Oi horizon. Arthropod abundance was best explained by forest floor mass, while arthropod diversity was best explained by phosphorus, calcium and sodium concentrations in the Oi horizon and by phosphorus concentrations in the Oe horizon. Differences in arthropod community composition between treatments and horizons correlated with phosphorus concentration and dry mass of the forest floor. We conclude that at a local scale, arthropod abundance is related to forest floor mass (habitat space), while arthropod diversity is related to forest floor nutrient concentrations (habitat quality).     

Reproductive Ecology of Understory Species in a Tropical Montane Forest in Costa Rica1

The reproductive ecology of nine hermaphroditic understory species in a tropical montane Quercus forest was studied at two sites (2300 and 2600 m elev.) in the Cordillera de Talamanca, Costa Rica. Flower life span, studied in six species, averaged 4.4 d. This is longer than flower life spans found in the Monteverde cloud forest (2.7 d) and comparable to flower life spans found for arctic and alpine species. We studied the breeding system in five species and found no self-incompatible species. Four species proved self-compatible, and three of these showed autogamy. The main diurnal insect pollinator was the bumblebee Bombus ephippiatus. Natural fruit set was low (8-32%) in six species with few seeds per fruit, while two many-seeded species showed a high rate of fruit set (90 and 96%). The incidence of pre–dispersal seed predation was high; the percentage of seeds infested in four species ranged from 8 to 56 percent.     

Pollinators of Syntopic Marcgravia Species in Costa Rican Lowland Rain Forest: Bats and Opossums

Abstract: The visitors of four syntopically occurring species of Marcgravia (M. nervosa, M. serrae, M. mexicana , and M. nepen-thoides ) in the Atlantic lowland rain forest at La Selva Biological Station in Costa Rica were observed. The four species differed in flowering phenology and morphology of inflorescences. Flowers of all species opened during the night and the stamens dropped before dawn. All species were visited by small nectar-feeding bats (Phyllostomidae: Glossophaginae). Marcgrovia nepen-thoides , with much larger nectaries, additionally attracted two species of opossums, Caluromys derbianus and Didelphis marsu-pious . Judging from the large distance between nectaries and flowers, glossophagine bats probably are inefficient pollinators of this species, and merely exploit a non-flying mammal pollination system.     

Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

Phenology, Lignotubers, and Water Relations of Cochlospermum vitifolium, a Pioneer Tropical Dry Forest Tree in Costa Rica

We examined structural and physiological traits relevant to the phenology of the tropical dry forest (TDF) pioneer tree Cochlospermum vitifolium . Despite marked seasonality in rainfall, meristem activity occurred throughout the year. Leaves were produced almost continuously during the rainy season, while leaf shedding started early during drought, before changes in soil water content were observed. Phenological activity under drought included flowering and fruiting of leafless trees; bud break and shoot extension took place before the end of the dry season. Low wood density of C. vitifolium stems (0.17 g/cm³) and lignotubers (0.14 g/cm³) provided water and starch storage needed to support phenological events such as branch extension, leaf flushing, and reproduction during the dry season, and probably also contributed to survival following mechanical damage and fire, typical of early TDF successional stages. Lignotuber water and starch contents showed substantial seasonal variation, declining from the beginning of the dry season to their lowest levels at the time of reproduction and dry-season flushing. Stems progressively replaced lignotubers as main storage organs as tree size increased. Evidence for a role of water stores in buffering daily water deficits was weak. Leaf water potentials remained above −1.2 MPa and stomatal conductance below 350 mmol/m²/s, suggesting that gas exchange during the rainy season was limited to prevent xylem cavitation. Leaf shedding occurred when early-morning and mid-day Ψ_L converged at the rainy–dry season transition, without changes in lignotuber or soil water content, suggesting that leaves of C. vitifolium are closely tuned to atmospheric drought.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .