One of the main environmental threats in the tropics is selective logging, which has degraded large areas of forest. In southeast Asia, enrichment planting with seedlings of the dominant group of dipterocarp tree species aims to accelerate restoration of forest structure and functioning. The role of tree diversity in forest restoration is still unclear, but the 'insurance hypothesis' predicts that in temporally and spatially varying environments planting mixtures may stabilize functioning owing to differences in species traits and ecologies. To test for potential insurance effects, we analyse the patterns of seedling mortality and growth in monoculture and mixture plots over the first decade of the Sabah biodiversity experiment. Our results reveal the species differences required for potential insurance effects including a trade-off in which species with denser wood have lower growth rates but higher survival. This trade-off was consistent over time during the first decade, but growth and mortality varied spatially across our 500 ha experiment with species responding to changing conditions in different ways. Overall, average survival rates were extreme in monocultures than mixtures consistent with a potential insurance effect in which monocultures of poorly surviving species risk recruitment failure, whereas monocultures of species with high survival have rates of self-thinning that are potentially wasteful when seedling stocks are limited. Longer-term monitoring as species interactions strengthen will be needed to more comprehensively test to what degree mixtures of species spread risk and use limited seedling stocks more efficiently to increase diversity and restore ecosystem structure and functioning.
Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.
Tree architecture, growth, and mortality change with increasing tree size and associated light conditions. To date, few studies have quantified how size-dependent changes in growth and mortality rates co-vary with architectural traits, and how such size-dependent changes differ across species and possible light capture strategies. We applied a hierarchical Bayesian model to quantify size-dependent changes in demographic rates and correlated demographic rates and architectural traits for 145 co-occurring Malaysian rain-forest tree species covering a wide range of tree sizes. Demographic rates were estimated using relative growth rate in stem diameter (RGR) and mortality rate as a function of stem diameter. Architectural traits examined were adult stature measured as the 95-percentile of the maximum stem diameter (upper diameter), wood density, and three tree architectural variables: tree height, foliage height, and crown width. Correlations between demographic rates and architectural traits were examined for stem diameters ranging from 1 to 47 cm. As a result, RGR and mortality varied significantly with increasing stem diameter across species. At smaller stem diameters, RGR was higher for tall trees with wide crowns, large upper diameter, and low wood density. Increased mortality was associated with low wood density at small diameters, and associated with small upper diameter and wide crowns over a wide range of stem diameters. Positive correlations between RGR and mortality were found over the whole range of stem diameters, but they were significant only at small stem diameters. Associations between architectural traits and demographic rates were strongest at small stem diameters. In the dark understory of tropical rain forests, the limiting amount of light is likely to make the interspecific difference in the effects of functional traits on demography more clear. Demographic performance is therefore tightly linked with architectural traits such as adult stature, wood density, and capacity for horizontal crown expansion. The enhancement of a demographic trade-off due to interspecific variation in functional traits in the understory helps to explain species coexistence in diverse rain forests.
Tropical beta diversity, and particularly that of herbivorous insects in rainforests, is often considered to be enormous, but this notion has recently been challenged. Because tropical beta diversity is highly relevant to our view on biodiversity, it is important to gain more insights and to resolve methodological problems that may lead to contradictions in different studies. We used data on two ecologically distinct moth families from Southeast Asia and analyzed separately the contribution of beta components to overall species richness at three spatial scales. Observed diversity partitions were compared under different types of null models. We found that alpha diversity was lower than expected on the basis of null models, whereas hierarchical beta components were larger than expected. Beta components played a significant role in shaping gamma diversity, and their contribution can be high (multiplicative beta >5). We found a reduction in beta components when comparing primary forests to agricultural sites (cf. "biotic homogenization"), but even in these habitats, beta components were still substantial. Our analyses show that beta components do play an important role in our data on tropical herbivorous insects and that these results are not attributable to lumping different habitats when sampling environmental gradients.
Diatom abundance, biovolume and diversity were measured over a 2-year period along the Straits of Malacca at two stations with upper (Klang) and lower (Port Dickson) states of eutrophication. Diatom abundance, which ranged from 0.2 × 10(4) to 21.7 × 10(4) cells L(-1) at Klang and 0.9 × 10(3)- 41.3 × 10(3) cells L(-1) at Port Dickson, was influenced partly by nutrient concentrations. At Klang, the diatoms were generally smaller and less diverse (H' = 0.77 ± 0.48) and predominated by Skeletonema spp. (60 ± 32% of total diatom biomass). In contrast, diatoms were larger and more diverse (H' = 1.40 ± 0.67) at Port Dickson. Chaetoceros spp. were the most abundant diatoms at Port Dickson but attributed only 48 ± 30% of total diatom biomass. Comparison of both Klang and Port Dickson showed that their diatom community structure differed and that eutrophication reduced diatom diversity at Klang. We also observed how Si(OH)4 affected the abundance of Skeletonema spp. which in turn influenced the temporal variation of diatom community at Klang. Our results highlighted how eutrophication affects diatom diversity and community structure.
Tree autotrophic respiratory processes, especially stem respiration or stem CO2 efflux (Estem), are important components of the forest carbon budget. Despite efforts to investigate the controlling processes of Estem in recent years, a considerable lack in our knowledge remains on the abiotic and biotic drivers affecting Estem dynamics. It has been strongly advocated that long-term measurements would shed light onto those processes. The expensive scientific instruments needed to measure gas exchange have prevented Estem measurements from being applied on a larger temporal and spatial scale. Here, we present an automated closed dynamic chamber system based on inexpensive and industrially broadly applied CO2 sensors, reducing the costs for the sensing system to a minimum. The CO2 sensor was cross-calibrated with a commonly used gas exchange system in the laboratory and in the field, and we found very good accordance of these sensors. We tested the system under harsh tropical climatic conditions, characterized by heavy tropical rainfall events, extreme humidity and temperatures, in a moist lowland forest in Malaysia. We recorded Estem of three Dyera costulata (Miq.) trees with our prototype over various days. The variation of Estem was large among the three tree individuals and varied by 7.5-fold. However, clear diurnal changes in Estem were present in all three tree individuals. One tree showed high diurnal variation in Estem, and the relationship between Estem and temperature was characterized by a strong hysteresis. The large variations found within one single tree species highlight the importance of continuous measurement to quantify ecosystem carbon fluxes.
Logging, pervasive across the lowland tropics, affects millions of hectares of forest, yet its influence on nutrient cycling remains poorly understood. One hypothesis is that logging influences phosphorus (P) cycling, because this scarce nutrient is removed in extracted timber and eroded soil, leading to shifts in ecosystem functioning and community composition. However, testing this is challenging because P varies within landscapes as a function of geology, topography and climate. Superimposed upon these trends are compositional changes in logged forests, with species with more acquisitive traits, characterized by higher foliar P concentrations, more dominant. It is difficult to resolve these patterns using traditional field approaches alone. Here, we use airborne light detection and ranging-guided hyperspectral imagery to map foliar nutrient (i.e. P, nitrogen [N]) concentrations, calibrated using field measured traits, over 400 km2 of northeastern Borneo, including a landscape-level disturbance gradient spanning old-growth to repeatedly logged forests. The maps reveal that canopy foliar P and N concentrations decrease with elevation. These relationships were not identified using traditional field measurements of leaf and soil nutrients. After controlling for topography, canopy foliar nutrient concentrations were lower in logged forest than in old-growth areas, reflecting decreased nutrient availability. However, foliar nutrient concentrations and specific leaf area were greatest in relatively short patches in logged areas, reflecting a shift in composition to pioneer species with acquisitive traits. N:P ratio increased in logged forest, suggesting reduced soil P availability through disturbance. Through the first landscape scale assessment of how functional leaf traits change in response to logging, we find that differences from old-growth forest become more pronounced as logged forests increase in stature over time, suggesting exacerbated phosphorus limitation as forests recover.
The existing knowledge regarding seawater intrusion and particularly upconing, in which both problems are linked to pumping, entirely relies on theoretical assumptions. Therefore, in this paper, an attempt is made to capture the effects of pumping on seawater intrusion and upconing using 2D resistivity measurement. For this work, two positions, one perpendicular and the other parallel to the sea, were chosen as profile line for resistivity measurement in the coastal area near the pumping wells of Kapas Island, Malaysia. Subsequently, water was pumped out of two pumping wells simultaneously for about five straight hours. Then, immediately after the pumping stopped, resistivity measurements were taken along the two stationed profile lines. This was followed by additional measurements after four and eight hours. The results showed an upconing with low resistivity of about 1-10 Ωm just beneath the pumping well along the first profile line that was taken just after the pumping stopped. The resistivity image also shows an intrusion of saline water (water enriched with diluted salt) from the sea coming towards the pumping well with resistivity values ranging between 10 and 25 Ωm. The subsequent measurements show the recovery of freshwater in the aquifer and how the saline water is gradually diluted or pushed out of the aquifer. Similarly the line parallel to the sea (L2) reveals almost the same result as the first line. However, in the second and third measurements, there were some significant variations which were contrary to the expectation that the freshwater may completely flush out the saline water from the aquifer. These two time series lines show that as the areas with the lowest resistivity (1 Ωm) shrink with time, the low resistivity (10 Ωm) tends to take over almost the entire area implying that the freshwater-saltwater equilibrium zone has already been altered. These results have clearly enhanced our current understanding and add more scientific weight to the theoretical assumptions on the effects of pumping on seawater intrusion and upconing.
Different mechanisms have been proposed to explain how vertical and horizontal heterogeneity in light conditions enhances tree species coexistence in forest ecosystems. The foliage partitioning theory proposes that differentiation in vertical foliage distribution, caused by an interspecific variation in mortality-to-growth ratio, promotes stable coexistence. In contrast, successional niche theory posits that horizontal light heterogeneity, caused by gap dynamics, enhances species coexistence through an interspecific trade-off between growth rate and survival. To distinguish between these theories of species coexistence, we analyzed tree inventory data for 370 species from the 50-ha plot in Pasoh Forest Reserve, Malaysia. We used community-wide Bayesian models to quantify size-dependent growth rate and mortality of every species. We compared the observed size distributions and the projected distributions from size-dependent demographic rates. We found that the observed size distributions were not simply correlated with the rate of population increase but were related to demographic properties such as size growth rate and mortality. Species with low relative abundance of juveniles in size distribution showed high growth rate and low mortality at small tree sizes and low per-capita recruitment rate. Overall, our findings were in accordance with those predicted by foliage partitioning theory.
Deforestation in the tropics is an important source of carbon C release to the atmosphere. To provide a sound scientific base for efforts taken to reduce emissions from deforestation and degradation (REDD+) good estimates of C stocks and fluxes are important. We present components of the C balance for selectively logged lowland tropical dipterocarp rainforest in the Malua Forest Reserve of Sabah, Malaysian Borneo. Total organic C in this area was 167.9 Mg C ha⁻¹±3.8 (SD), including: Total aboveground (TAGC: 55%; 91.9 Mg C ha⁻¹±2.9 SEM) and belowground carbon in trees (TBGC: 10%; 16.5 Mg C ha⁻¹±0.5 SEM), deadwood (8%; 13.2 Mg C ha⁻¹±3.5 SEM) and soil organic matter (SOM: 24%; 39.6 Mg C ha⁻¹±0.9 SEM), understory vegetation (3%; 5.1 Mg C ha⁻¹±1.7 SEM), standing litter (<1%; 0.7 Mg C ha⁻¹±0.1 SEM) and fine root biomass (<1%; 0.9 Mg C ha⁻¹±0.1 SEM). Fluxes included litterfall, a proxy for leaf net primary productivity (4.9 Mg C ha⁻¹ yr⁻¹±0.1 SEM), and soil respiration, a measure for heterotrophic ecosystem respiration (28.6 Mg C ha⁻¹ yr⁻¹±1.2 SEM). The missing estimates necessary to close the C balance are wood net primary productivity and autotrophic respiration.Twenty-two years after logging TAGC stocks were 28% lower compared to unlogged forest (128 Mg C ha⁻¹±13.4 SEM); a combined weighted average mean reduction due to selective logging of -57.8 Mg C ha⁻¹ (with 95% CI -75.5 to -40.2). Based on the findings we conclude that selective logging decreased the dipterocarp stock by 55-66%. Silvicultural treatments may have the potential to accelerate the recovery of dipterocarp C stocks to pre-logging levels.
With a focus on the Danum Valley area of Sabah, Malaysian Borneo, this special issue has as its theme the future of tropical rainforests in a changing landscape and climate. The global environmental context to the issue is briefly given before the contents and rationale of the issue are summarized. Most of the papers are based on research carried out as part of the Royal Society South East Asia Rainforest Research Programme. The issue is divided into five sections: (i) the historical land-use and land management context; (ii) implications of land-use change for atmospheric chemistry and climate change; (iii) impacts of logging, forest fragmentation (particularly within an oil palm plantation landscape) and forest restoration on ecosystems and their functioning; (iv) the response and resilience of rainforest systems to climatic and land-use change; and (v) the scientific messages and policy implications arising from the research findings presented in the issue.
The characteristic flavor of exotic tropical fruits is one of their most attractive attributes to consumers. In this article, the enormous diversity of exotic fruit flavors is reviewed. Classifying some of the exotic fruits into two classes on the basis of whether esters or terpenes predominate in the aroma was also attempted. Indeed, as far as exotic tropical fruits are concerned, the majority of fruits have terpenes predominating in their aroma profile. Some of the fruits in this group are the Amazonian fruits such as pitanga, umbu-caja, camu-camu, garcinia, and bacuri. The ester group is made up of rambutan, durians, star fruit, snake fruit, acerola, tamarind, sapodilla, genipap, soursop, cashew, melon, jackfruit, and cupuacu respectively. Also, the role of sulphur-volatiles in some of the exotic fruits is detailed.
This paper examines the storm runoff quality from a commercial area in south Johor, Malaysia. Six storm events with a total of 68 storm runoff samples were analyzed. Event Mean Concentration (EMC) for all constituents analysed showed large inter-event variation. Site mean concentrations (SMC) for total suspended solids (TSS), oil and grease (O&G), biochemical oxygen demand (BOD), chemical oxygen demand (COD), nitrate-nitrogen (NO(3)-N), nitrite-nitrogen (NO(2)-N), ammonia-nitrogen (NH(3)-N), total phosphorus (Total P) and Soluble P are 261, 4.31, 74, 192, 1.5, 0.006, 1.9, 1.12 and 0.38 mg/L, respectively. The SMCs at the studied site are higher than those reported in many urban catchments. The mean baseflow concentrations were higher than the EMCs for COD, Soluble P, NH(3)-N, NO(3)-N, Total P and NO(2)-N. However, the reverse was observed for TSS and O&G. All pollutants showed the occurrence of first flush phenomenon with the highest strength was observed for TSS, COD and NH(3)-N.
Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.
This article presents findings of the thermal comfort study in hospitals. A field survey was conducted to investigate the temperature range for thermal comfort in hospitals in the tropics. Thermal acceptability assessment was conducted to examine whether the hospitals in the tropics met the ASHRAE Standard-55 80% acceptability criteria. A total of 114 occupants in four hospitals were involved in the study. The results of the field study revealed that only 44% of the examined locations met the comfort criteria specified in ASHRAE Standard 55. The survey also examined the predicted percentage of dissatisfied in the hospitals. The results showed that 49% of the occupants were satisfied with the thermal environments in the hospitals. The field survey analysis revealed that the neutral temperature for Malaysian hospitals was 26.4 degrees C. The comfort temperature range that satisfied 90% of the occupants in the space was in the range of 25.3-28.2 degrees C. The results from the field study suggested that a higher comfort temperature was required for Malaysians in hospital environments compared with the temperature criteria specified in ASHRAE Standard (2003). In addition, the significant deviation between actual mean vote and predicted mean vote (PMV) strongly implied that PMV could not be applied without errors in hospitals in the tropics.
There is accumulating evidence that similar suites of plant traits may affect leaf palatability and leaf litter decomposability. However, the possible association between leaf herbivory and litter decomposition rates across species in species-diverse natural ecosystems such as tropical rain forests remains unexplored, despite its importance in estimating the herbivory effects on carbon and nutrient cycling of ecosystems. We found no strong association between leaf herbivory and litter decomposition rates across 40 tree species in a Malaysian tropical rain forest, even though the leaf and litter traits were tightly correlated. This is because the leaf and litter traits related to herbivory and decomposition rates in the field were inconsistent. Leaf toughness accounted for only a small part of the variation in the herbivory rate, whereas a number of litter traits (the leaf mass per area, lignin to nitrogen ratio, and condensed tannin concentration) accurately predicted the decomposition rate across species. These results suggest that herbivory rate across species may not be strongly related to single leaf traits, probably because plant-herbivore interactions in tropical rain forests are highly diverse; on the other hand, plant-decomposer interactions are less specific and can be governed by litter chemicals. We also investigated two factors, phylogeny and tree functional types, that could affect the relationship between herbivory and decomposition across species. Phylogenetic relatedness among the species did not affect the relationship between herbivory and decomposition. In contrast, when the plants were segregated according to their leaf emergence pattern, we found a significant positive relationship between herbivory and decomposition rates for continuous-leafing species. In these species, the condensed tannin to N ratios in leaves and litter were related to herbivory and decomposition rates, respectively. However, we did not observe a similar trend for synchronous-leafing species. These results suggest that the relationship between herbivory and decomposition may be more greatly affected by functional types than by phylogenetic relatedness among species. In conclusion, our results suggest that well-defended leaves are not necessarily less decomposable litter in a tropical rain forest community, implying that herbivory may not generate positive feedback for carbon and nutrient cycling in this type of ecosystem.
The responses of tropical forests to global anthropogenic disturbances remain poorly understood. Above-ground woody biomass in some tropical forest plots has increased over the past several decades, potentially reflecting a widespread response to increased resource availability, for example, due to elevated atmospheric CO2 and/or nutrient deposition. However, previous studies of biomass dynamics have not accounted for natural patterns of disturbance and gap phase regeneration, making it difficult to quantify the importance of environmental changes. Using spatially explicit census data from large (50 ha) inventory plots, we investigated the influence of gap phase processes on the biomass dynamics of four 'old-growth' tropical forests (Barro Colorado Island (BCI), Panama; Pasoh and Lambir, Malaysia; and Huai Kha Khaeng (HKK), Thailand). We show that biomass increases were gradual and concentrated in earlier-phase forest patches, while biomass losses were generally of greater magnitude but concentrated in rarer later-phase patches. We then estimate the rate of biomass change at each site independent of gap phase dynamics using reduced major axis regressions and ANCOVA tests. Above-ground woody biomass increased significantly at Pasoh (+0.72% yr(-1)) and decreased at HKK (-0.56% yr(-1)) independent of changes in gap phase but remained stable at both BCI and Lambir. We conclude that gap phase processes play an important role in the biomass dynamics of tropical forests, and that quantifying the role of gap phase processes will help improve our understanding of the factors driving changes in forest biomass as well as their place in the global carbon budget.
A comparative study of understorey birds inhabiting different habitats, that is, virgin jungle reserve (VJR) and regenerated forest (RF), was conducted in Ulu Gombak Forest Reserve and Selangor and Triang Forest Reserve, Negeri Sembilan, Peninsular Malaysia. The objective of this study was to assess the diversity of understorey birds in both habitats and the effects of forest regeneration on the understorey bird community. The mist-netting method was used to capture understorey birds inhabiting both habitats in both locations. Species composition and feeding guild indicated that understorey bird populations were similar in the two habitats. However, the number of secondary forest species such as Little spiderhunter (Arachnothera longirostra) in VJR is increasing due to its proximity to RF. This study discovered that RFs in both study areas are not yet fully recovered. However, based on the range of species discovered, the RFs have conservation value and should be maintained because they harbour important forest species such as babblers and flycatchers. The assessment of the community structure of understorey birds in VJR and RF is important for forest management and conservation, especially where both habitats are intact.