The Malaysian states of Sabah and Sarawak are global hotspots of forest loss and degradation due to timber and oil palm industries; however, the rates and patterns of change have remained poorly measured by conventional field or satellite approaches. Using 30 m resolution optical imagery acquired since 1990, forest cover and logging roads were mapped throughout Malaysian Borneo and Brunei using the Carnegie Landsat Analysis System. We uncovered ∼364,000 km of roads constructed through the forests of this region. We estimated that in 2009 there were at most 45,400 km(2) of intact forest ecosystems in Malaysian Borneo and Brunei. Critically, we found that nearly 80% of the land surface of Sabah and Sarawak was impacted by previously undocumented, high-impact logging or clearing operations from 1990 to 2009. This contrasted strongly with neighbouring Brunei, where 54% of the land area remained covered by unlogged forest. Overall, only 8% and 3% of land area in Sabah and Sarawak, respectively, was covered by intact forests under designated protected areas. Our assessment shows that very few forest ecosystems remain intact in Sabah or Sarawak, but that Brunei, by largely excluding industrial logging from its borders, has been comparatively successful in protecting its forests.
Although a substantial body of evidence suggests that large and old trees have reduced metabolic levels, the search for the causes behind this observation has proved elusive. The strong coupling between age and size, commonly encountered in the field, precludes the isolation of the potential causes. We used standard propagation techniques (grafting and air-layering) to decouple the effects of size from those of age in affecting leaf structure, biochemistry and physiology of two broadleaved trees, Acer pseudoplatanus (a diffuse-porous species) and Fraxinus excelsior (a ring-porous species). The first year after establishment of the propagated plants, some of the measurements suggested the presence of age-related declines in metabolism, while other measurements either did not show any difference or suggested variability across treatments not associated with either age or size. During the second year after establishment, only one of the measured properties (specific leaf area) continued to show some evidence of an age-mediated decline (although much reduced compared to the field), whereas, for some properties (particularly for F. excelsior), even the opposite trend of age-related increases was apparent. We concluded that (1) our plants suffered from grafting shock during year 1 and they gradually recovered during year 2; (2) the results over 2 years do not support the statement that age directly mediates ageing in either species but instead suggest that size directly mediates ageing processes; and (3) neither shoots nor roots of A. pseudoplatanus showed any evidence of senescence.
The carnivorous Utricularia (Lentibulariaceae) is a small herb of multifarious wet habitats worldwide. Eleven of the 14 Peninsular Malaysian species range into the mountains. Distribution, disturbance adaptability and collection frequency were used to formulate their commonness category. Common (U. aurea, U. bifida, and U. minutissima) and fairly common (U. gibba and U. uliginosa) species are mostly lowland plants that ascend to open montane microhabitats, while the fairly common (U. striatula), narrow-range (U. caerulea pink form and U. involvens), rare (U. furcellata and U. scandens), and endemic (U. vitellina) species are restricted to mountainous sites. Common species that colonise dystrophic to oligotrophic man-made sites in late succession could serve as predictors for general health and recovery of wet habitats. Rarer species are often locally abundant, their niches situated around pristine forest edges. When in decline, they indicate the beginning of problems affecting the forest. Utricularia is reportedly nutritious, mildly astringent, and diuretic. Preadapted to nutrient-poor, waterlogged soils, U. bifida is suitable as an alternative for small-scale herb cultivation on low pH, wet poor soils usually deemed not suitable for any crops.
Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.
In South-East Asian dipterocarp forests, many trees synchronize their reproduction at the community level, but irregularly, in a phenomenon known as general flowering (GF). Several proximate cues have been proposed as triggers for the synchronization of Southeast Asian GF, but the debate continues, as many studies have not considered geographical variation in climate and flora. We hypothesized that the spatial pattern of GF forests is explained by previously proposed climatic cues if there are common cues for GF among regions. During the study, GF episodes occurred every year, but the spatial occurrence varied considerably from just a few forests to the whole of Peninsular Malaysia. In 2001, 2002 and 2005, minor and major GF occurred widely throughout Peninsular Malaysia (GF2001, GF2002, and GF2005), and the geographical patterns of GF varied between the episodes. In the three regional-scale GF episodes, most major events occurred in regions where prolonged drought (PD) had been recorded prior, and significant associations between GF scores and PD were found in GF2001 and GF2002. However, the frequency of PD was higher than that of GF throughout the peninsula. In contrast, low temperature (LT) was observed during the study period only before GF2002 and GF2005, but there was no clear spatial relationship between GF and LT in the regional-scale episodes. There was also no evidence that last GF condition influenced the magnitude of GF. Thus, our results suggest that PD would be essential to trigger regional-scale GF in the peninsula, but also that PD does not fully explain the spatial and temporal patterns of GF. The coarse relationships between GF and the proposed climatic cues may be due to the geographical variation in proximate cues for GF, and the climatic and floristic geographical variations should be considered to understand the proximate factors of GF.
Tropical deforestation is occurring at an alarming rate, threatening the ecological integrity of protected areas. This makes it vital to regularly assess protected areas to confirm the efficacy of measures that protect that area from clearing. Satellite remote sensing offers a systematic and objective means for detecting and monitoring deforestation. This paper examines a spectral change approach to detect deforestation using pattern decomposition (PD) coefficients from multitemporal Landsat data. Our results show that the PD coefficients for soil and vegetation can be used to detect deforestation using change vector analysis (CVA). CVA analysis demonstrates that deforestation in the Kinabalu area, Sabah, Malaysia has significantly slowed from 1.2% in period 1 (1973 and 1991) to 0.1% in period 2 (1991 and 1996). A comparison of deforestation both inside and outside Kinabalu Park has highlighted the effectiveness of the park in protecting the tropical forest against clearing. However, the park is still facing pressure from the area immediately surrounding the park (the 1 km buffer zone) where the deforestation rate has remained unchanged.
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.
Partitioning of soil phosphorus (P) pools has been proposed as a key mechanism maintaining plant diversity, but experimental support is lacking. Here, we provided different chemical forms of P to 15 tree species with contrasting root symbiotic relationships to investigate plant P acquisition in both tropical and subtropical forests. Both ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) trees responded positively to addition of inorganic P, but strikingly, ECM trees acquired more P from a complex organic form (phytic acid). Most ECM tree species and all AM tree species also showed some capacity to take up simple organic P (monophosphate). Mycorrhizal colonisation was negatively correlated with soil extractable P concentration, suggesting that mycorrhizal fungi may regulate organic P acquisition among tree species. Our results support the hypothesis that ECM and AM plants partition soil P sources, which may play an ecologically important role in promoting species coexistence in tropical and subtropical forests.
The impacts of global change on tropical forests remain poorly understood. We examined changes in tree growth rates over the past two decades for all species occurring in large (50-ha) forest dynamics plots in Panama and Malaysia. Stem growth rates declined significantly at both forests regardless of initial size or organizational level (species, community or stand). Decreasing growth rates were widespread, occurring in 24-71% of species at Barro Colorado Island, Panama (BCI) and in 58-95% of species at Pasoh, Malaysia (depending on the sizes of stems included). Changes in growth were not consistently associated with initial growth rate, adult stature, or wood density. Changes in growth were significantly associated with regional climate changes: at both sites growth was negatively correlated with annual mean daily minimum temperatures, and at BCI growth was positively correlated with annual precipitation and number of rainfree days (a measure of relative insolation). While the underlying cause(s) of decelerating growth is still unresolved, these patterns strongly contradict the hypothesized pantropical increase in tree growth rates caused by carbon fertilization. Decelerating tree growth will have important economic and environmental implications.
Among the local processes that determine species diversity in ecological communities, fluctuation-dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.
Accurate estimation of tree biomass is necessary to provide realistic values of the carbon stored in the terrestrial biosphere. A recognized source of errors in tree aboveground biomass (AGB) estimation is introduced when individual tree height values (H) are not directly measured but estimated from diameter at breast height (DBH) using allometric equations. In this paper, we evaluate the performance of 12 alternative DBH : H equations and compare their effects on AGB estimation for three tropical forests that occur in contrasting climatic and altitudinal zones. We found that fitting a three-parameter Weibull function using data collected locally generated the lowest errors and bias in H estimation, and that equations fitted to these data were more accurate than equations with parameters derived from the literature. For computing AGB, the introduced error values differed notably among DBH : H allometric equations, and in most cases showed a clear bias that resulted in either over- or under-estimation of AGB. Fitting the three-parameter Weibull function minimized errors in AGB estimates in our study and we recommend its widespread adoption for carbon stock estimation. We conclude that many previous studies are likely to present biased estimates of AGB due to the method of H estimation.
Periods of extreme food abundance, such as irregular masting events, can dramatically affect animal populations and communities, but the extent to which anthropogenic disturbances alter animal responses to mast events is not clear. In South-East Asia, dipterocarp trees reproduce in mast fruiting events every 2-10 years in some of the largest masting events on the planet. These trees, however, are targeted for selective logging, reducing the intensity of fruit production and potentially affecting multiple trophic levels. Moreover, animal responses to resource pulse events have largely been studied in systems where the major mast consumers have been extirpated. We sought to evaluate the influence of human-induced habitat disturbance on animal responses to masting in a system where key mast consumers remain extant. We used motion-triggered camera traps to quantify terrestrial mammal and bird occurrences in Sabah, Malaysian Borneo, relative to variation in fruit biomass from 69 plant families during a major (2014) and minor (2015) masting event and a non-mast year (2013), in both logged and unlogged forests. Bearded pigs (Sus barbatus) showed the clearest responses to masting and occurrence rates were highest in unlogged forest in the year following the major mast, suggesting that the pulse in fruit availability increased immigration or reproduction. We also detected local-scale spatial tracking of dipterocarp fruits in bearded pigs in unlogged forest, while this was equivocal in other species. In contrast, pigs and other vertebrate taxa in our study showed limited response to spatial or temporal variation in fruit availability in logged forest. Our findings suggest that vertebrates, namely bearded pigs, may respond to masting via movement and increased reproduction, but that these responses may be attenuated by habitat disturbance.
Successful cost-effective reforestation plantings depend substantially on maximising sapling survival from the time of planting, yet in reforestation programs remarkably little attention is given to management of saplings at the planting stage and to planting methods used. Critical determinants of sapling survival include their vigour and condition when planted, the wetness of the soil into which saplings are planted, the trauma of transplant shock from nursery to natural field soils, and the method and care taken during planting. While some determinants are outside planters' control, careful management of specific elements associated with outplanting can significantly lessen transplanting shock and improve survival rates. Results from three reforestation experiments designed to examine cost-effective planting methods in the Australian wet tropics provided the opportunity to examine the effects of specific planting treatments, including (1) watering regime prior to planting, (2) method of planting and planter technique, and (3) site preparation and maintenance, on sapling survival and establishment. Focusing on sapling root moisture and physical protection during planting improved sapling survival by at least 10% (>91% versus 81%) at 4 months. Survival rates of saplings under different planting treatments were reflected in longer-term survival of trees at 18-20 months, differing from a low of 52% up to 76-88%. This survival effect was evident more than 6 years after planting. Watering saplings immediately prior to planting, careful planting using a forester's planting spade in moist soil and suppressing grass competition using appropriate herbicides were critical to improved plant survival.
Tree species of Eurasian broadleaved forest possess two divergent trait syndromes with contrasting patterns of resource allocation adapted to different selection environments: short-stature basal resprouters that divert resources to a bud bank adapted to frequent and severe disturbances such as fire and herbivory, and tall trees that delay reproduction by investing in rapid height growth to escape shading. Drawing on theory developed in savanna ecosystems, we propose a conceptual framework showing that the possession of contrasting trait syndromes is essential for the persistence of broadleaved trees in an open ecosystem that burns. Consistent with this hypothesis, trees of modern Eurasian broadleaved forest bear a suite of traits that are adaptive to surface and crown-fire regimes. We contend that limited opportunities in grassland restricts recruitment to disturbance-free refugia, and en masse establishment creates a wooded environment where shade limits the growth of light-demanding savanna plants. Rapid height growth, which involves investment in structural support and the switch from a multi-stemmed to a monopodial growth form, is adaptive in this shaded environment. Although clustering reduces surface fuel loads, these establishment nuclei are vulnerable to high-intensity crown fires. The lethal effects of canopy fire are avoided by seasonal leaf shedding, and aerial resprouting enhances rapid post-fire recovery of photosynthetic capacity. While these woody formations satisfy the structural definition of forest, their constituents are clearly derived from savanna. Contrasting trait syndromes thus represent the shift from consumer to resource regulation in savanna ecosystems. Consistent with global trends, the diversification of most contemporary broadleaved taxa coincided with the spread of grasslands, a surge in fire activity and a decline in wooded ecosystems in the late Miocene-Pliocene. Recognition that Eurasian broadleaved forest has savanna origins and persists as an alternative state with adjacent grassy ecosystems has far-reaching management implications in accordance with functional rather than structural criteria. Shade is a severe constraint to the regeneration and growth of both woody and herbaceous growth forms in consumer-regulated ecosystems. However, these ecosystems are highly resilient to disturbance, an essential process that maintains diversity especially among the species-rich herbaceous component that is vulnerable to shading when consumer behaviour is altered.
Accurate estimates of forest biomass stocks and fluxes are needed to quantify global carbon budgets and assess the response of forests to climate change. However, most forest inventories consider tree mortality as the only aboveground biomass (AGB) loss without accounting for losses via damage to living trees: branchfall, trunk breakage, and wood decay. Here, we use ~151,000 annual records of tree survival and structural completeness to compare AGB loss via damage to living trees to total AGB loss (mortality + damage) in seven tropical forests widely distributed across environmental conditions. We find that 42% (3.62 Mg ha-1 year-1 ; 95% confidence interval [CI] 2.36-5.25) of total AGB loss (8.72 Mg ha-1 year-1 ; CI 5.57-12.86) is due to damage to living trees. Total AGB loss was highly variable among forests, but these differences were mainly caused by site variability in damage-related AGB losses rather than by mortality-related AGB losses. We show that conventional forest inventories overestimate stand-level AGB stocks by 4% (1%-17% range across forests) because assume structurally complete trees, underestimate total AGB loss by 29% (6%-57% range across forests) due to overlooked damage-related AGB losses, and overestimate AGB loss via mortality by 22% (7%-80% range across forests) because of the assumption that trees are undamaged before dying. Our results indicate that forest carbon fluxes are higher than previously thought. Damage on living trees is an underappreciated component of the forest carbon cycle that is likely to become even more important as the frequency and severity of forest disturbances increase.
The role of conspecific density dependence (CDD) in the maintenance of species richness is a central focus of tropical forest ecology. However, tests of CDD often ignore the integrated effects of CDD over multiple life stages and their long-term impacts on population demography. We combined a 10-year time series of seed production, seedling recruitment and sapling and tree demography of three dominant Southeast Asian tree species that adopt a mast-fruiting phenology. We used these data to construct individual-based models that examine the effects of CDD on population growth rates (λ) across life-history stages. Recruitment was driven by positive CDD for all species, supporting the predator satiation hypothesis, while negative CDD affected seedling and sapling growth of two species, significantly reducing λ. This negative CDD on juvenile growth overshadowed the positive CDD of recruitment, suggesting the cumulative effects of CDD during seedling and sapling development has greater importance than the positive CDD during infrequent masting events. Overall, CDD varied among positive, neutral and negative effects across life-history stages for all species, suggesting that assessments of CDD on transitions between just two stages (e.g. seeds seedlings or juveniles mature trees) probably misrepresent the importance of CDD on population growth and stability.
Differences in the density of conspecific tree individuals in response to environmental gradients are well documented for many tree species, but how such density differences are generated and maintained is poorly understood. We examined the segregation of six dipterocarp species among three soil types in the Pasoh tropical forest, Malaysia. We examined how individual performance and population dynamics changed across the soil types using 10-year demographic data to compare tree performance across soil types, and constructed population matrix models to analyze the population dynamics. Species showed only minor changes in mortality and juvenile growth across soil types, although recruitment differed greatly. Clear, interspecific demographic trade-offs between growth and mortality were found in all soil types. The relative trade-offs by a species did not differ substantially among the soil types. Population sizes were projected to remain stable in all soil types for all species with one exception. Our life-table response experiment demonstrated that the population dynamics of a species differed only subtly among soil types. Therefore, species with strong density differences across soil types do not necessarily differ greatly in their population dynamics across the soil types. In contrast, interspecific differences in population dynamics were large. The trade-off between mortality and growth led to a negative correlation between the contributions of mortality and growth to variations in the population growth rate (λ) and thus reduced their net contributions. Recruitment had little impact on the variation in λ. The combination of these factors resulted in little variation in λ among species.
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.
Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.
The formation of protein homodimer complexes for molecular catalysis and regulation is fascinating. The homodimer formation through 2S (2 state), 3SMI (3 state with monomer intermediate) and 3SDI (3 state with dimer intermediate) folding mechanism is known for 47 homodimer structures. Our dataset of forty-seven homodimers consists of twenty-eight 2S, twelve 3SMI and seven 3SDI. The dataset is characterized using monomer length, interface area and interface/total (I/T) residue ratio. It is found that 2S are often small in size with large I/T ratio and 3SDI are frequently large in size with small I/T ratio. Nonetheless, 3SMI have a mixture of these features. Hence, we used these parameters to develop a decision tree model. The decision tree model produced positive predictive values (PPV) of 72% for 2S, 58% for 3SMI and 57% for 3SDI in cross validation. Thus, the method finds application in assigning homodimers with folding mechanism.