Tropical rainforests are subject to extensive degradation by commercial selective logging. Despite pervasive changes to forest structure, selectively logged forests represent vital refugia for global biodiversity. The ability of these forests to buffer temperature-sensitive species from climate warming will be an important determinant of their future conservation value, although this topic remains largely unexplored. Thermal buffering potential is broadly determined by: (i) the difference between the "macroclimate" (climate at a local scale, m to ha) and the "microclimate" (climate at a fine-scale, mm to m, that is distinct from the macroclimate); (ii) thermal stability of microclimates (e.g. variation in daily temperatures); and (iii) the availability of microclimates to organisms. We compared these metrics in undisturbed primary forest and intensively logged forest on Borneo, using thermal images to capture cool microclimates on the surface of the forest floor, and information from dataloggers placed inside deadwood, tree holes and leaf litter. Although major differences in forest structure remained 9-12 years after repeated selective logging, we found that logging activity had very little effect on thermal buffering, in terms of macroclimate and microclimate temperatures, and the overall availability of microclimates. For 1°C warming in the macroclimate, temperature inside deadwood, tree holes and leaf litter warmed slightly more in primary forest than in logged forest, but the effect amounted to <0.1°C difference between forest types. We therefore conclude that selectively logged forests are similar to primary forests in their potential for thermal buffering, and subsequent ability to retain temperature-sensitive species under climate change. Selectively logged forests can play a crucial role in the long-term maintenance of global biodiversity.
Flies attracted to human remains during death investigations were surveyed in north Peninsular Malaysia. Six families, eight genera, and 16 species were identified from human remains, with the greatest fly diversity occurring on remains recovered indoors. The total relative frequency of species was led by Chrysomya megacephala (Fabricius, 1794) (46%), followed by Chrysomya rufifacies (Macquart, 1842) (22%), Sarcophaga (Liopygia) ruficornis (Fabricius, 1974) (5%), Sarcophaga spp. (4%), Synthesiomyia nudiseta Wulp, 1883 (6%), Megaselia spp. (3%), Megaselia scalaris (Loew, 1866), (2%), Megaselia spiracularis Schmitz, 1938 (2%), and Chrysomya villeneuvi Patton, 1922 (2%). Hemipyrellia tagaliana (Bigot, 1877), Desmometopa sp., Megaselia curtineura (Brues, 1909), Hemipyrellia ligurriens Wiedemann 1830, Ophyra sp., Sarcophaga princeps Wiedemann 1830, Piophila casei (Linnaeus, 1758), and unidentified pupae each represented 1%, respectively.
The extent to which environmental heterogeneity can account for tree species coexistence in diverse ecosystems, such as tropical rainforests, is hotly debated, although the importance of spatial variability in contributing to species co-existence is well recognized. Termites contribute to the micro-topographical and nutrient spatial heterogeneity of tropical forests. We therefore investigated whether epigeal termite mounds could contribute to the coexistence of plant species within a 50 ha plot at Pasoh Forest Reserve, Malaysia. Overall, stem density was significantly higher on mounds than in their immediate surroundings, but tree species diversity was significantly lower. Canonical correspondence analysis showed that location on or off mounds significantly influenced species distribution when stems were characterized by basal area. Like studies of termite mounds in other ecosystems, our results suggest that epigeal termite mounds provide a specific microhabitat for the enhanced growth and survival of certain species in these species-rich tropical forests. However, the extent to which epigeal termite mounds facilitate species coexistence warrants further investigation.
We examined relationships between mortality rate, relative growth rate (RGR), and spatial patterns of three growth stages (small, medium, and large trees) for 11 dipterocarp species in the Pasoh 50-ha plot. Mortality rates for these species tended to be positively correlated with RGRs, although the correlation was significant only at the small-tree stage. Seven species with high growth and mortality rates exhibited peaks in spatial aggregation at small distances (<100 m) in small trees, but this aggregation disappeared in medium and large trees. In contrast, the other four species with low growth and mortality rates aggregated at large distances (>200 m) throughout the three growth stages in all but one species. Negative associations between different growth stages were observed only for the high-mortality species, suggesting density-dependent mortality. The high-mortality species showed habitat associations with topography, soil type, and the forest regeneration phase after gap formation, whereas the three low-mortality species only had associations with the forest regeneration phase. A randomization procedure revealed that these habitat associations explained little of their spatial aggregation. Our results suggest that the growth strategy has a large effect on the structuring of the spatial distribution of tree species through mortality processes.
Mesohabitat selection in fluvial fishes was studied in a small tropical stream of the Malay Peninsula. A total of 681 individuals representing 24 species were sampled at 45 stations within heterogeneous stream (ca. 1 km in length), in which water depth, water velocity, substrate size, and riparian canopy cover were measured as environmental variables. A canonical correspondence analysis (CCA) yielded a diagram that shows a specific mesohabitat selection of the fish assemblage, in which the species were plotted widely on the CCA1-CCA2 biplot. Generalized linear model also revealed a significant pattern of the mesohabitat selection of several species. Water velocity and substrate size mainly separated on CCA1, indicating variation of pool (deep, slow-flow section) and riffle (shallow, fast-flow section) structures is a primary factor of mesohabitat selection in the fluvial fish assemblage. The mean body weight of species significantly correlated with CCA1; larger species tended to inhabit pools, while small ones occupied riffles. The riparian canopy cover separated on CCA2. The trophic level of species significantly correlated with CCA2; herbivorous species (low trophic level) selected open sites without riparian cover, whereas omnivorous/carnivorous (middle-high trophic level) species preferred highly covered sites. In conclusion, our results suggest that mesohabitat selection is closely related to the species feeding habit, which is consistent with the results of previous studies.
A power law called the species-area relationship describes the finding that the number of species is proportional to the size of the area in which they are found, raised to an exponent (usually, a number between 0.2 and 0.3). In their Perspective, May and Stumpf discuss new results from a survey of five tropical forest census areas containing a total of a million trees. They explain how this large data set can be used to fine-tune the existing power law so that it provides a better prediction of species diversity in small census samples.
The response of tropical forests to global climate variability and change remains poorly understood. Results from long-term studies of permanent forest plots have reported different, and in some cases opposing trends in tropical forest dynamics. In this study, we examined changes in tree growth rates at four long-term permanent tropical forest research plots in relation to variation in solar radiation, temperature and precipitation. Temporal variation in the stand-level growth rates measured at five-year intervals was found to be positively correlated with variation in incoming solar radiation and negatively related to temporal variation in night-time temperatures. Taken alone, neither solar radiation variability nor the effects of night-time temperatures can account for the observed temporal variation in tree growth rates across sites, but when considered together, these two climate variables account for most of the observed temporal variability in tree growth rates. Further analysis indicates that the stand-level response is primarily driven by the responses of smaller-sized trees (less than 20 cm in diameter). The combined temperature and radiation responses identified in this study provide a potential explanation for the conflicting patterns in tree growth rates found in previous studies.
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.
Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.
Pristine tropical peat swamp forests (PSFs) represent a unique wetland ecosystem of distinctive hydrology which support unique biodiversity and globally significant stores of soil carbon. Yet in Indonesia and Malaysia, home to 56% of the world's tropical peatland, they are subject to considerable developmental pressures, including widespread drainage to support agricultural needs. In this article, we review the ecology behind the functioning and ecosystem services provided by PSFs, with a particular focus on hydrological processes as well as the role of the forest itself in maintaining those services. Drawing on this, we review the suitability of current policy frameworks and consider the efficacy of their implementation. We suggest that policies in Malaysia and Indonesia are often based around the narrative of oil palm and other major monocrops as drivers of prosperity and development. However, we also argue that this narrative is also being supported by a priori claims concerning the possibility of sustainability of peat swamp exploitation via drainage-based agriculture through the adherence to best management practices. We discuss how this limits their efficacy, uptake and the political will towards enforcement. Further, we consider how both narratives (prosperity and sustainability) clearly exclude important considerations concerning the ecosystem value of tropical PSFs which are dependent on their unimpacted hydrology. Current research clearly shows that the actual debate should be focused not on how to develop drainage-based plantations sustainably, but on whether the sustainable conversion to drainage-based systems is possible at all.
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.
Outdoor mass culture of microalgae in the tropical area is important to minimize its production cost. This study evaluates the growth of Chaetoceros calcitrans in 120 L annular photobioreactors at indoor temperature (Treatment I, 25 +/- 2 degrees C) and outdoor tropical ambient temperature, (Treatment II, 30 +/- 6 degrees C). Each treatment was done in duplicates. For both treatments, C. calcitrans was first grown in starter columns of 10 L capacity for a period of 7 days at 25 +/- 2 degrees C. After 7 days, the 9 L culture was transferred to the annular photobioreactors and subsequently brought to a final volume of 100 L by adding 20 L fresh medium every 5 days. There was no significant difference (p > 0.05) in the dry weight of microalgae grown in natural light and those grown indoor. The results suggest that C. calcitrans can be grown in outdoor conditions, hence, saving time and microalgae production cost for the larviculture industry.
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.
Inadequate funding from developed countries has hampered international efforts to conserve biodiversity in tropical forests. We present two complementary research approaches that reveal a significant increase in public demand for conservation within tropical developing countries as those countries reach upper-middle-income (UMI) status. We highlight UMI tropical countries because they contain nearly four-fifths of tropical primary forests, which are rich in biodiversity and stored carbon. The first approach is a set of statistical analyses of various cross-country conservation indicators, which suggests that protective government policies have lagged behind the increase in public demand in these countries. The second approach is a case study from Malaysia, which reveals in a more integrated fashion the linkages from rising household income to increased household willingness to pay for conservation, nongovernmental organization activity, and delayed government action. Our findings suggest that domestic funding in UMI tropical countries can play a larger role in (i) closing the funding gap for tropical forest conservation, and (ii) paying for supplementary conservation actions linked to international payments for reduced greenhouse gas emissions from deforestation and forest degradation in tropical countries.
Differences in demographic and environmental niches facilitate plant species coexistence in tropical forests. However, the adaptations that enable species to achieve higher demographic rates (e.g. growth or survival) or occupy unique environmental niches (e.g. waterlogged conditions) remain poorly understood. Anatomical traits may better predict plant environmental and demographic strategies because they are direct measurements of structures involved in these adaptations. We collected 18 leaf and twig traits from 29 tree species in a tropical freshwater swamp forest in Singapore. We estimated demographic parameters of the 29 species from growth and survival models, and degree of association toward swamp habitats. We examined pairwise trait-trait, trait-demography and trait-environment links while controlling for phylogeny. Leaf and twig anatomical traits were better predictors of all demographic parameters than other commonly measured leaf and wood traits. Plants with wider vessels had faster growth rates but lower survival rates. Leaf and spongy mesophyll thickness predicted swamp association. These findings demonstrate the utility of anatomical traits as indicators of plant hydraulic strategies and their links to growth-mortality trade-offs and waterlogging stress tolerance that underlie species coexistence mechanisms in tropical forest trees.
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.
The data presented in this paper is supporting the research article "Life history traits predict the response to increased light among 33 tropical rainforest tree species" [3]. We show basic growth and survival data collected over the 6 years duration of the experiment, as well as data from traits inventories covering 12 tree traits collected prior to and after a canopy reduction treatment in 2013. Further, we also include canopy closure and forest light environment data from measurements with hemispherical photographs before and after the treatment.
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 application of pultruded glass fiber-reinforced polymer composites (PGFRPCs) as a replacement for conventional wooden cross-arms in transmission towers is relatively new. Although numerous studies have conducted creep tests on coupon-scale PGFRPC cross-arms, none had performed creep analyses on full-scale PGFRPC cross-arms under actual working load conditions. Thus, this work proposed to study the influence of an additional bracing system on the creep responses of PGFRPC cross-arms in a 132 kV transmission tower. The creep behaviors and responses of the main members in current and braced PGFRPC cross-arm designs were compared and evaluated in a transmission tower under actual working conditions. These PGFRPC cross-arms were subjected to actual working loads mimicking the actual weight of electrical cables and insulators for a duration of 1000 h. The cross-arms were installed on a custom test rig in an open area to simulate the actual environment of tropical climate conditions. Further creep analysis was performed by using Findley and Burger models on the basis of experimental data to link instantaneous and extended (transient and viscoelastic) creep strains. The addition of braced arms to the structure reduced the total strain of a cross-arm's main member beams and improved elastic and viscous moduli. The addition of bracing arms improved the structural integrity and stiffness of the cross-arm structure. The findings of this study suggested that the use of a bracing system in cross-arm structures could prolong the structures' service life and subsequently reduce maintenance effort and cost for long-term applications in transmission towers.