Rahayu Sukmaria Sukri

Ecological implications of a flower size/number trade-off in tropical forest trees

Background In angiosperms, flower size commonly scales negatively with number. The ecological consequences of this trade-off for tropical trees remain poorly resolved, despite their potential importance for tropical forest conservation. We investigated the flower size number trade-off and its implications for fecundity in a sample of tree species from the Dipterocarpaceae on Borneo. Methodology/Principal Findings We combined experimental exclusion of pollinators in 11 species, with direct and indirect estimates of contemporary pollen dispersal in two study species and published estimates of pollen dispersal in a further three species to explore the relationship between flower size, pollinator size and mean pollen dispersal distance. Maximum flower production was two orders of magnitude greater in small-flowered than large-flowered species of Dipterocarpaceae. In contrast, fruit production was unrelated to flower size and did not differ significantly among species. Small-flowered species had both smaller-sized pollinators and lower mean pollination success than large-flowered species. Average pollen dispersal distances were lower and frequency of mating between related individuals was higher in a smaller-flowered species than a larger-flowered confamilial. Our synthesis of pollen dispersal estimates across five species of dipterocarp suggests that pollen dispersal scales positively with flower size. Conclusions and Their Significance Trade-offs embedded in the relationship between flower size and pollination success contribute to a reduction in the variance of fecundity among species. It is therefore plausible that these processes could delay competitive exclusion and contribute to maintenance of species coexistence in this ecologically and economically important family of tropical trees. These results have practical implications for tree species conservation and restoration. Seed collection from small-flowered species may be especially vulnerable to cryptic genetic erosion. Our findings also highlight the potential for differential vulnerability of tropical tree species to the deleterious consequences of forest fragmentation.

Distinctive Tropical Forest Variants Have Unique Soil Microbial Communities, But Not Always Low Microbial Diversity

There has been little study of whether different variants of tropical rainforest have distinct soil microbial communities and levels of diversity. We compared bacterial and fungal community composition and diversity between primary mixed dipterocarp, secondary mixed dipterocarp, white sand heath, inland heath, and peat swamp forests in Brunei Darussalam, Northwest Borneo by analyzing Illumina Miseq sequence data of 16S rRNA gene and ITS1 region. We hypothesized that white sand heath, inland heath and peat swamp forests would show lower microbial diversity and relatively distinct microbial communities (compared to MDF primary and secondary forests) due to their distinctive environments. We found that soil properties together with bacterial and fungal communities varied significantly between forest types. Alpha and beta-diversity of bacteria was highest in secondary dipterocarp and white sand heath forests. Also, bacterial alpha diversity was strongly structured by pH, adding another instance of this widespread pattern in nature. The alpha diversity of fungi was equally high in all forest types except peat swamp forest, although fungal beta-diversity was highest in primary and secondary mixed dipterocarp forests. The relative abundance of ectomycorrhizal (EcM) fungi varied significantly between forest types, with highest relative abundance observed in MDF primary forest. Overall, our results suggest that the soil bacterial and fungal communities in these forest types are to a certain extent predictable and structured by soil properties, but that diversity is not determined by how distinctive the conditions are. This contrasts with the diversity patterns seen in rainforest trees, where distinctive soil conditions have consistently lower tree diversity.

Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edge effects

Less than half of anthropogenic carbon dioxide emissions remain in the atmosphere. While carbon balance models imply large carbon uptake in tropical forests, direct on-the-ground observations are still lacking in Southeast Asia. Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 per year (95% CI 0.14–0.72, mean period 1988–2010) in above-ground live biomass carbon. These results closely match those from African and Amazonian plot networks, suggesting that the world’s remaining intact tropical forests are now en masse out-of-equilibrium. Although both pan-tropical and long-term, the sink in remaining intact forests appears vulnerable to climate and land use changes. Across Borneo the 1997–1998 El Niño drought temporarily halted the carbon sink by increasing tree mortality, while fragmentation persistently offset the sink and turned many edge-affected forests into a carbon source to the atmosphere.The existence of a pan-tropical forest carbon sink remains uncertain due to the lack of data from Asia. Here, using direct on-the-ground observations, the authors confirm remaining intact forests in Borneo have provided a long-term carbon sink, but carbon net gains are vulnerable to drought and edge effects.

Tree Diversity and Community Composition of the Tutong White Sands, Brunei Darussalam: A Rare Tropical Heath Forest Ecosystem

Bornean heath (Kerangas) forests are a unique and increasingly rare tropical forest ecosystem that remains little studied. We quantified tree floristic diversity in Kerangas forests in the Tutong White Sands, Brunei Darussalam, and investigated the influence of soil and environmental variables on community composition. Six 20 m × 20 m plots were established, where all trees of ≥5 cm diameter at breast height (DBH) were identified and measured to determine stem diameter and basal area. We determined pH, gravimetric water content, and concentrations of total nitrogen (N) and phosphorus (P) in topsoil, as well as litter depth and percentage canopy openness. A total of 296 trees were recorded, representing 78 species in 59 genera and 38 families. Stem diameter, basal area, species richness, and species diversity differed significantly among the six plots. The NMDS ordination revealed that differences in tree community compositions were significantly associated with total N concentrations and percentage canopy openness. Despite the small sampling area, we recorded several Bornean endemic tree species (16/78 tree species; 20.5%), including several IUCN Red List endangered and vulnerable species. Our results illustrate the potentially high conservation value of the Kerangas forests in the Tutong White Sands and highlight the urgent need to protect and conserve this area.

Thermochemical characterisation of Acacia auriculiformis tree parts via proximate, ultimate, TGA, DTG, calorific value and FTIR spectroscopy analyses to evaluate their potential as a biofuel resource

ABSTRACT Continuously increasing energy requirements coupled with environmental pollution have established pressure to utilise lignocellulosic biomass for energy production. Acacia auriculiformis is a fast-growing species capable of accumulating large quantities of biomass without requiring major agricultural inputs. The aim of this research was to investigate the thermochemical properties of its tree parts including phyllodes (leaves), trunk, bark and branches to utilise them as solid fuel to produce bioenergy. Thermogravimetric and derivative thermogravimetric (TGA and DTG ) analyses were performed to study the biomass degradation behaviour, which showed the decomposition of biomass in three major stages corresponding to the decomposition of hemicellulose, cellulose and lignin components. Fourier transform infrared (FTIR) analysis was carried out to determine the functional groups. Proximate analysis showed the weight percentages of moisture contents, volatile matter, fixed carbon and ash contents as 7.25–9.27%, 61.79–73.28%, 16.50–27.92% and 2.13–3.72%, respectively. Ultimate analysis showed the ranges of carbon, hydrogen and oxygen as 44.27–49.41%, 5.3–6.10% and 41.93–49.44% respectively, while lower values of sulphur and nitrogen components were reported which are encouraging from an environmental perspective. Higher heating values (HHV) for the parts were reported to range between 17.85 and 20.93 MJ/kg on a dry basis.

Field methods for sampling tree height for tropical forest biomass estimation

Abstract Quantifying the relationship between tree diameter and height is a key component of efforts to estimate biomass and carbon stocks in tropical forests. Although substantial site‐to‐site variation in height–diameter allometries has been documented, the time consuming nature of measuring all tree heights in an inventory plot means that most studies do not include height, or else use generic pan‐tropical or regional allometric equations to estimate height. Using a pan‐tropical dataset of 73 plots where at least 150 trees had in‐field ground‐based height measurements, we examined how the number of trees sampled affects the performance of locally derived height–diameter allometries, and evaluated the performance of different methods for sampling trees for height measurement. Using cross‐validation, we found that allometries constructed with just 20 locally measured values could often predict tree height with lower error than regional or climate‐based allometries (mean reduction in prediction error = 0.46 m). The predictive performance of locally derived allometries improved with sample size, but with diminishing returns in performance gains when more than 40 trees were sampled. Estimates of stand‐level biomass produced using local allometries to estimate tree height show no over‐ or under‐estimation bias when compared with biomass estimates using field measured heights. We evaluated five strategies to sample trees for height measurement, and found that sampling strategies that included measuring the heights of the ten largest diameter trees in a plot outperformed (in terms of resulting in local height–diameter models with low height prediction error) entirely random or diameter size‐class stratified approaches. Our results indicate that even limited sampling of heights can be used to refine height–diameter allometries. We recommend aiming for a conservative threshold of sampling 50 trees per location for height measurement, and including the ten trees with the largest diameter in this sample.

Height-diameter input data and R-code to fit and assess height-diameter models, from 'Field methods for sampling tree height for tropical forest biomass estimation' in Methods in Ecology and Evolution

1. Quantifying the relationship between tree diameter and height is a key component of efforts to estimate biomass and carbon stocks in tropical forests. Although substantial site-to-site variation in height-diameter allometries has been documented, the time consuming nature of measuring all tree heights in an inventory plot means that most studies do not include height, or else use generic pan-tropical or regional allometric equations to estimate height. 2. Using a pan-tropical dataset of 73 plots where at least 150 trees had in-field ground-based height measurements, we examined how the number of trees sampled affects the performance of locally-derived height-diameter allometries, and evaluated the performance of different methods for sampling trees for height measurement. 3. Using cross-validation, we found that allometries constructed with just 20 locally measured values could often predict tree height with lower error than regional or climate-based allometries (mean reduction in prediction error = 0.46 m). The predictive performance of locally-derived allometries improved with sample size, but with diminishing returns in performance gains when more than 40 trees were sampled. Estimates of stand-level biomass produced using local allometries to estimate tree height show no over- or under-estimation bias when compared with estimates using measured heights. We evaluated five strategies to sample trees for height measurement, and found that sampling strategies that included measuring the heights of the ten largest diameter trees in a plot outperformed (in terms of resulting in local height-diameter models with low height prediction error) entirely random or diameter size-class stratified approaches. 4. Our results indicate that even remarkably limited sampling of heights can be used to refine height-diameter allometries. We recommend aiming for a conservative threshold of sampling 50 trees per location for height measurement, and including the ten trees with the largest diameter in this sample.

Author Correction: Long-term carbon sink in Borneo’s forests halted by drought and vulnerable to edge effects

The original version of this Article contained an error in the third sentence of the abstract and incorrectly read “Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 year−1 (95% CI 0.14–0.72, mean period 1988–2010) above-ground live biomass”, rather than the correct “Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha−1 year−1 (95% CI 0.14–0.72, mean period 1988–2010) in above-ground live biomass carbon”. This has now been corrected in both the PDF and HTML versions of the Article.

Distinctive soil archaeal communities in different variants of tropical equatorial forest

Little is known of how soil archaeal community composition and diversity differ between local variants of tropical rainforests. We hypothesized that (1) as with plants, animals, fungi, and bacteria, the soil archaeal community would differ between different variants of tropical forest; (2) that spatially rarer forest variants would have a less diverse archaeal community than common ones; (3) that a history of forest disturbance would decrease archaeal alpha- and beta-diversity; and (4) that archaeal distributions within the forest would be governed more by deterministic than stochastic factors. We sampled soil across several different forest types within Brunei, Northwest Borneo. Soil DNA was extracted, and the 16S rRNA gene of archaea was sequenced using Illumina MiSeq. We found that (1) as hypothesized, there are distinct archaeal communities for each forest type, and community composition significantly correlates with soil parameters including pH, organic matter, and available phosphorous. (2) As hypothesized, the “rare” white sand forest variants kerangas and inland heath had lower archaeal diversity. A nestedness analysis showed that archaeal community in inland heath and kerangas was mainly a less diverse subset of that in dipterocarp forests. However, primary dipterocarp forest had the lowest beta-diversity among the other tropical forest types. (3) Also, as predicted, forest disturbance resulted in lower archaeal alpha-diversity—but increased beta-diversity in contrast with our predictions. (4) Contrary to our predictions, the BetaNTI of the various primary forest types indicated community assembly was mainly stochastic. The possible effects of these habitat and disturbance-related effects on N cycling should be investigated.

Photosynthesis in Co-occurring Invasive Acacia spp. and Native Bornean Heath Forest Trees at the Post-establishment Invasion Stage

ABSTRACT Seedlings of invasive species often exhibit superior physiological traits that facilitate their spread at early stages of invasion, although it is unclear whether these traits persist at the post-establishment stage. To determine whether mature exotic Acacia spp. possess superior traits over mature native plants, we compared foliar gas exchange and chlorophyll a fluorescence of Acacia auriculiformis and Acacia mangium coexisting with tropical heath forest tree species Buchanania arborescens and Dillenia suffruticosa in Brunei Darussalam. The CO2 assimilation rates of Acacia spp. were significantly higher than those of heath species at current prevailing conditions of ~400 ppm ambient CO2, 1,500 µmol m−2 s−1 photosynthetically active radiation and 30°C leaf temperature. The photosystem II of Acacia spp. exhibited significantly higher maximum quantum yield of primary photochemistry at comparable temperatures, and was more sensitive to an elevated temperature (42°C for 1 h). Better photosynthetic performance of Acacia spp., due to larger stomatal openings, better light harvesting efficiency, and greater plasticity in photosystem II, may enable adult Acacia trees to sustain a competitive growth advantage and suppress native tropical heath forest species. The competitive advantage maintained by Acacia spp. in post-establishment stage likely facilitates the establishment of monospecific Acacia stands in invaded heath forests.