The exceptionally high species richness of arthropods in tropical rainforests hinges on the complexity of the forest itself: that is, on features such as the high plant diversity, the layered nature of the canopy and the abundance and the diversity of epiphytes and litter. We here report on one important, but almost completely neglected, piece of this complex jigsaw-the intricate network of rhizomorph-forming fungi that ramify through the vegetation of the lower canopy and intercept falling leaf litter. We show that this litter-trapping network is abundant and intercepts substantial amounts of litter (257.3 kg ha(-1)): this exceeds the amount of material recorded in any other rainforest litter-trapping system. Experimental removal of this fungal network resulted in a dramatic reduction in both the abundance (decreased by 70.2 ± 4.1%) and morphospecies richness (decreased by 57.4 ± 5.1%) of arthropods. Since the lower canopy levels can contain the highest densities of arthropods, the proportion of the rainforest fauna dependent on the fungal networks is likely to be substantial. Fungal litter-trapping systems are therefore a crucial component of habitat complexity, providing a vital resource that contributes significantly to rainforest biodiversity.
The diversity and abundance of macroinvertebrate shredders were investigated in 52 forested streams (local scale) from nine catchments (regional scale) covering a large area of peninsular Malaysia. A total of 10,642 individuals of aquatic macroinvertebrates were collected, of which 18.22% were shredders. Biodiversity of shredders was described by alpha (αaverage), beta (β) and gamma diversity (γ) measures. We found high diversity and abundance of shredders in all catchments, represented by 1,939 individuals (range 6-115 and average per site of 37.29±3.48 SE) from 31 taxa with 2-13 taxa per site (αaverage=6.98±0.33 SE) and 10-15 taxa per catchment (γ=13.33±0.55 SE). At the local scale, water temperature, stream width, depth and altitude were correlated significantly with diversity (Adj-R2=0.205). Meanwhile, dissolved oxygen, stream velocity, water temperature, stream width and altitude were correlated to shredder abundance (Adj-R2=0.242). At regional scale, however, water temperature was correlated negatively with β and γ diversity (r2=0.161 and 0.237, respectively) as well as abundance of shredders (r2=0.235). Canopy cover was correlated positively with β diversity (r2=0.378) and abundance (r2=0.266), meanwhile altitude was correlated positively with β (quadratic: r2=0.175), γ diversity (quadratic: r2=0.848) as well as abundance (quadratic: r2=0.299). The present study is considered as the first report describing the biodiversity and abundance of shredders in forested headwater streams across a large spatial scale in peninsular Malaysia. We concluded that water temperature has a negative effect while altitude showed a positive relationship with diversity and abundance of shredders. However, it was difficult to detect an influence of canopy cover on shredder diversity.
We monitored the reproductive status of all trees with diameters at breast height (dbh) >30 cm in a 40-ha plot at Pasoh, west Malaysia, and investigated the individual fecundity of 15 Shorea acuminata Dyer (Dipterocarpaceae) trees using seed-trapping methods during two consecutive general flowering periods in 2001 (GF2001) and 2002 (GF2002). The proportion of flowering trees was higher, and not dependent on size, in GF2002 (84.2%), than in GF2001 (54.5%), when flowering mainly occurred in trees with a dbh < or =70 cm. Fecundity parameters of individual trees per event varied widely (221,000-35,200,000 flowers, 0-139,000 mature seeds, and 1.04-177 kg total dry matter mass of fruit (TDM) per tree). Monotonic increases with increasing tree size were observed for flower production and TDM amongst trees up to 90 cm in dbh, but not for mature seed production or for any of these parameters amongst larger trees. The pattern of reproductive investment during the two consecutive reproductive events clearly differed between medium-sized and large trees; the former concentrated their reproductive investment in one of the reproductive events whereas the latter allocated their investment more evenly to both reproductive events. Our results suggest size-related differences in the resource allocation pattern for reproduction.
Dispersal-assembly theories of species coexistence posit that environmental factors play no role in explaining community diversity and structure. Dispersal-assembly theories shed light on some aspects of community structure such as species-area and species-abundance relationships. However, species' environmental associations also affect these measures of community structure. Measurements of species' niche breadth and overlap address this influence. Using a new continuous measure of niche and a dispersal-assembly null model that maintains species' niche breadth and aggregation, we tested two hypotheses assessing the effects of habitat heterogeneity on the ability of dispersal-assembly theories to explain community niche structure. We found that in both homogenous and heterogeneous environments dispersal-assembly theories cannot fully explain observed niche structure. The performance of the dispersal-assembly null models was particularly poor in heterogeneous environments. These results indicate that non-dispersal based mechanisms are in part responsible for observed community structure and measures of community structure which include species' environmental associations should be used to test theories of species diversity.
The effects of selective logging on the diversity and species composition of moths were investigated by sampling from multiple sites in primary forest, both understorey and canopy, and logged forest at Danum Valley, Sabah, Malaysia. The diversity of individual sites was similar, although rarefied species richness of logged forest was 17% lower than for primary forest (understorey and canopy combined). There was significant heterogeneity in faunal composition and measures of similarity (NESS index) among primary forest understorey sites which may be as great as those between primary understorey and logged forest. The lowest similarity values were between primary forest understorey and canopy, indicating a distinct canopy fauna. A number of species encountered in the logged forest were confined to, or more abundant in, the canopy of primary forest. Approximately 10% of species were confined to primary forest across a range of species' abundances, suggesting this is a minimum estimate for the number of species lost following logging. The importance of accounting for heterogeneity within primary forest and sampling in the canopy when measuring the effects of disturbance on tropical forest communities are emphasized.
Leaf samples of tropical trees, i.e. Dryobalanops lanceolata (Kapur paji), Dipterocarpaceae and Macaranga spp. (Mahang), Euphorbiaceae were analyzed for 21 chemical elements. The pioneer Macaranga spp. exhibited higher concentrations for the majority of elements compared to the emergent species of Dryobalanops lanceolata, which was attributed to the higher physiological activity of the fast growing pioneer species compared to emergent trees. Lead showed rather high concentrations in several samples from the Bakam re-forestation site. This is suggested to be caused by emissions through brick manufacturing and related activities in the vicinity. A comparison of Dryobalanops lanceolata samples collected in 1993, 1995 and 1997 in the Lambir Hills National Park revealed that certain heavy metals, i.e. Co, Cu, Mn, Ni, Pb and Ti showed higher values in 1997 compared to the previous years, which could indicate an atmospheric input from the haze caused by the extensive forest fires raging in Borneo and other parts of Southeast Asia.
The behavioral response of the obligate bamboo-nesting ant Cataulacus muticus to nest flooding was studied in a perhumid tropical rainforest in Malaysia and in the laboratory. The hollow internodes of giant bamboo, in which C. muticus exclusively nests, are prone to flooding by heavy rains. The ants showed a two-graded response to flooding. During heavy rain workers block the nest entrances with their heads to reduce water influx. However, rainwater may still intrude into the nest chamber. The ants respond by drinking the water, leaving the nest and excreting water droplets on the outer stem surface. This cooperative 'peeing' behavior is a new survival mechanism adaptive to the ants' nesting ecology. Laboratory experiments conducted with two other Cataulacus species, C. catuvolcus colonizing small dead twigs and C. horridus inhabiting rotten wood, did not reveal any form of water-bailing behavior.
Pollen flow and population genetic structure among 30 potentially flowering individuals of Neobalanocarpus heimii, a tropical emergent tree, were investigated in a lowland tropical rainforest of Malaysia using microsatellite polymorphism. The 248 offspring in the vicinity of five reproductive trees of the 30 potentially flowering trees were used in paternity analysis for pollen-flow study. Four primer pairs, developed in different species of dipterocarps, were adopted to detect microsatellite polymorphism. Based upon microsatellite polymorphism, pollen flow and seed migration were detected. Pollen-flow events of more than 400 m were observed directly, based on paternity analysis in the study plot. The estimated average mating distance of the five reproductive trees was 524 m. This result suggests that reproduction of this species is mediated by a long-distance pollinator. The haplotypes of some offspring were not compatible with the nearest reproductive tree. Thus, the results suggest that some seeds are dispersed by a seed dispersal vector. Investigation of genetic structure showed significant and negative correlation of genetic relatedness and spatial distances between the 30 potentially flowering trees, but this correlation was weak. We suggest that long-distance gene flow and seed migration are responsible for the poorly developed genetic structure of this species.
Quadrat-based analysis of two rainforest plots of area 50 ha, one in Panama (Barro Colorado Island, BCI) and the other in Malaysia (Pasoh), shows that in both plots recruitment is in general negatively correlated with both numbers and biomass of adult trees of the same species in the same quadrat. At BCI, this effect is not significantly influenced by treefall gaps. In both plots, recruitment of individual species is negatively correlated with the numbers of trees of all species in the quadrats, but not with overall biomass. These observations suggest, but do not prove, widespread frequency-dependent effects produced by pathogens and seed-predators that act most effectively in quadrats crowded with trees. Within-species correlations of mortality with numbers or biomass are not found in either plot, indicating that most frequency-dependent mortality takes place before the trees reach 1 cm in diameter. Stochastic effects caused by BCI's more rapid tree turnover may contribute to a larger variance in diversity from quadrat to quadrat at BCI, although they are not sufficient to explain why BCI has fewer than half as many tree species as Pasoh. Finally, in both plots quadrats with low diversity show a significant increase in diversity over time, and this increase is stronger at BCI. This process, like the frequency-dependence, will tend to maintain diversity over time. In general, these non-random forces that should lead to the maintenance of diversity are slightly stronger at BCI, even though the BCI plot is less diverse than the Pasoh plot.
Fully mapped tree census plots of large area, 25 to 52 hectares, have now been completed at six different sites in tropical forests, including dry deciduous to wet evergreen forest on two continents. One of the main goals of these plots has been to evaluate spatial patterns in tropical tree populations. Here the degree of aggregation in the distribution of 1768 tree species is examined based on the average density of conspecific trees in circular neighborhoods around each tree. When all individuals larger than 1 centimeter in stem diameter were included, nearly every species was more aggregated than a random distribution. Considering only larger trees (>/= 10 centimeters in diameter), the pattern persisted, with most species being more aggregated than random. Rare species were more aggregated than common species. All six forests were very similar in all the particulars of these results.
Data are presented from a long-term study of banded langurs in three contrasting rain forest habitats in Peninsular Malaysia. Results from different sites and months are used to correlate ranging patterns with food availability and other environmental variables. Day range lengths are correlated with availability of preferred foods; the degree of territoriality is related to the distribution and size of food sources and length of time for which any one of these produces favoured food items.
Precipitation patterns are changing across the globe causing more severe and frequent drought for many forest ecosystems. Although research has focused on the resistance of tree populations and communities to these novel precipitation regimes, resilience of forests is also contingent on recovery following drought, which remains poorly understood, especially in aseasonal tropical forests. We used rainfall exclusion shelters to manipulate the interannual frequency of drought for diverse seedling communities in a tropical forest and assessed resistance, recovery and resilience of seedling growth and mortality relative to everwet conditions. We found seedlings exposed to recurrent periods of drought altered their growth rates throughout the year relative to seedlings in everwet conditions. During drought periods, seedlings grew slower than seedlings in everwet conditions (i.e., resistance phase) while compensating with faster growth after drought (i.e., recovery phase). However, the response to frequent drought was species dependent as some species grew significantly slower with frequent drought relative to everwet conditions while others grew faster with frequent drought due to overcompensating growth during the recovery phase. In contrast, mortality was unrelated to rainfall conditions and instead correlated with differences in light. Intra-annual plasticity of growth and increased annual growth of some species led to an overall maintenance of growth rates of tropical seedling communities in response to more frequent drought. These results suggest these communities can potentially adapt to predicted climate change scenarios and that plasticity in the growth of species, and not solely changes in mortality rates among species, may contribute to shifts in community composition under drought.
Salmacis sphaeroides (Linnaeus, 1758) is one of the regular echinoids, occuring in the warm Indo-West Pacific, including Johor Straits, between Malaysia and Singapore. In order to investigate the developmental basis of morphological changes in embryos and larvae, we documented the ontogeny of S. sphaeroides in laboratory condition. Gametes were obtained from adult individuals by 0.5 M KCl injection into the coelomic cavity. Fertilization rate at limited sperm concentration (10(-5) dilution) was 96.6 ± 1.4% and the resulting embryos were reared at 24°C. First cleavage (2-cell), 4-cell, 8-cell, 16-cell, 32-cell, and multicell (Morulla) stages were achieved 01.12, 02.03, 02.28, 02.51, 03.12, and 03.32 h postfertilization. Ciliated blastulae with a mean length of 174.72 ± 4.43 μm hatched 08.45 h after sperm entry. The gastrulae formed 16.15 h postfertilization and the archenteron elongated constantly while ectodermal red-pigmented cells migrated synchronously to the apical plate. Pluteus larva started to feed unicellular algae in 2 d, grew continuously, and finally attained metamorphic competence in 35 d after fertilization. Metamorphosis took approximately 1 h 30 min from attachment to the complete resorption of larval tissues and the development of complete juvenile structure with adult spines, extended tubefeet and well-developed pedicellaria, the whole event of which usually took place within 1 d postsettlement. This study represents the first successful investigation on embryonic, larval, and early juvenile development of S. sphaeroides. The findings would greatly be helpful towards the understanding of ontogeny and life-history strategies, which will facilitate us to develop the breeding, seed production, and culture techniques of sea urchins in captive condition.
Intensive aeration for nitrification is a major energy consumer in sewage treatment plants (STPs). Low-dissolved-oxygen (low-DO) nitrification has the potential to lower the aeration demand. However, the applicability of low-DO nitrification in the tropical climate is not well-understood. In this study, the potential of low-DO nitrification in tropical setting was first examined using batch kinetic experiments. Subsequently, the performance of low-DO nitrification was investigated in a laboratory-scale sequential batch reactor (SBR) for 42 days using real tropical sewage. The batch kinetic experiments showed that the seed sludge has a relatively high oxygen affinity. Thus, the rate of nitrification was not significantly reduced at low DO concentrations (0.5 mg/L). During the operation of the low-DO nitrification SBR, 90% of NH4-N was removed. The active low-DO nitrification was mainly attributed to the limited biodegradable organics in the sewage. Fluorescence in-situ hybridisation and 16S rRNA amplicon sequencing revealed the nitrifiers were related to Nitrospira genus and Nitrosomonadaceae family. Phylogenetic analysis suggests 47% of the operational taxonomic units in Nitrospira genus are closely related to a comammox bacteria. This study has demonstrated active low-DO nitrification in tropical setting, which is a more sustainable process that could significantly reduce the energy footprint of STPs.
The associations among yield-related traits and the pattern of influence on rice grain yield were investigated. This evaluation is important to determine the direct and indirect effects of various traits on yield to determine selection criteria for higher grain yield. Fifteen rice genotypes were evaluated under tropical condition at five locations in two planting seasons. The experiment was laid out in a randomized complete block design with three replications across the locations. Data were collected on vegetative and yield components traits. The pooled data based on the analysis of variance revealed that there were significant differences (p < 0.001) among the fifteen genotypes for all the characters studied except for panicle length and 100-grain weight. Highly significant and positive correlations at phenotypic level were observed in grain weight per hill (0.796), filled grains per panicle (0.702), panicles per hill (0.632), and tillers per hill (0.712) with yield per hectare, while moderate positive correlations were observed in flag leaf length to width ratio (0.348), days to flowering (0.412), and days to maturity (0.544). By contrast, unfilled grains per panicle (-0.225) and plant height (-0.342) had a negative significant association with yield per hectare. Filled grains per panicle (0.491) exhibited the maximum positive direct effect on yield followed by grain weight per hill (0.449), while unfilled grain per panicle (-0.144) had a negative direct effect. The maximum indirect effect on yield per hectare was recorded by the tillers per hill through the panicles per hill. Therefore, tillers per hill, filled grains per panicle, and grain weight per hill could be used as selection criteria for improving grain yield in rice.
Both niche and stochastic dispersal processes structure the extraordinary diversity of tropical plants, but determining their relative contributions has proven challenging. We address this question using airborne imaging spectroscopy to estimate canopy β-diversity for an extensive region of a Bornean rainforest and challenge these data with models incorporating niches and dispersal. We show that remotely sensed and field-derived estimates of pairwise dissimilarity in community composition are closely matched, proving the applicability of imaging spectroscopy to provide β-diversity data for entire landscapes of over 1000 ha containing contrasting forest types. Our model reproduces the empirical data well and shows that the ecological processes maintaining tropical forest diversity are scale dependent. Patterns of β-diversity are shaped by stochastic dispersal processes acting locally whilst environmental processes act over a wider range of scales.
Tree autotrophic respiratory processes, especially stem respiration or stem CO2 efflux (Estem), are important components of the forest carbon budget. Despite the efforts to investigate the controlling processes of Estem in the last 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 into those processes. The expensive scientific instruments needed to measure gas exchange has prevented from applying Estem measurements 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 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 highlights the importance of continuous measurement to quantify ecosystem carbon fluxes.