Effects of land use on surface-atmosphere exchanges of trace gases and energy in Borneo: comparing fluxes over oil palm plantations and a rainforest

This paper reports measurements of land-atmosphere fluxes of sensible and latent heat, momentum, CO(2), volatile organic compounds (VOCs), NO, NO(2), N(2)O and O(3) over a 30 m high rainforest canopy and a 12 m high oil palm plantation in the same region of Sabah in Borneo between April and July 2008. The daytime maximum CO(2) flux to the two canopies differs by approximately a factor of 2, 1200 mg C m(-2) h(-1) for the oil palm and 700 mg C m(-2) h(-1) for the rainforest, with the oil palm plantation showing a substantially greater quantum efficiency. Total VOC emissions are also larger over the oil palm than over the rainforest by a factor of 3. Emissions of isoprene from the oil palm canopy represented 80 per cent of the VOC emissions and exceeded those over the rainforest in similar light and temperature conditions by on average a factor of 5. Substantial emissions of estragole (1-allyl-4-methoxybenzene) from the oil palm plantation were detected and no trace of this VOC was detected in or above the rainforest. Deposition velocities for O(3) to the rainforest were a factor of 2 larger than over oil palm. Emissions of nitrous oxide were larger from the soils of the oil palm plantation than from the soils of the rainforest by approximately 25 per cent. It is clear from the measurements that the large change in the species composition generated by replacing rainforest with oil palm leads to profound changes in the net exchange of most of the trace gases measured, and thus on the chemical composition of the boundary layer over these surfaces.     

Ecological impacts of oil palm agriculture on forest mammals in plantation estates and smallholdings

Global oil palm expansion has caused substantial ecological damage to tropical biodiversity. We quantified wild mammal richness in large oil palm plantation estates and semi-traditional oil palm smallholdings in Peninsular Malaysia. We sampled 41 plantation estates and 14 smallholdings, and used line-transect surveys coupled with semi-structured interviews to develop a database of the native mammals found in oil palm landscapes. Semi-structured interviews revealed a total of 32 mammal species, including 13 IUCN Red Listed taxa of high conservation value. Our results showed that human activity and the size of patches of remnant rainforest were important factors influencing the richness of mammal species in oil palm landscapes. More carnivorous and herbivorous species were reported in smallholdings than plantation estates, most probably as a response to greater habitat heterogeneity in smallholdings. All species, irrespective of conservation status, were more likely to be recorded in oil palm plantation estates and smallholdings that supported large areas of native forest. Our findings suggest that biodiversity conservation in oil palm landscapes will require a variety of conservation approaches. Minimizing poaching, reducing disturbance from human activity, and protecting existing forest patches appear particularly important. Strategies to promote the persistence of both high and low conservation value species should be adopted as part of a strengthened certification scheme for oil palm production.     

环境因子对植物释放挥发性化合物的影响

对近年来有关环境因子与植物释放挥发性化合物关系的研究进展进行了综合和概括。本文主要包括3类挥发性化合物。⑴异戊二烯是由叶绿体产生并且直接释放到大气中的C5化合物。⑵单萜类化合物是一类环状或非环状的C10化合物,它在植物体内合成后首先贮存于体内的特殊结构中（如树脂道、油腺）,然后由此通过气孔向大气中释放。⑶含氧挥发性化合物以各种形式释放大大气中。它包括醇、醛、酮、酯和有机酸。本文的重点是前两者,主要阐述了二方面内容：⑴植物军发性化合物的生物合成和释放机理。⑵环境因子（如温度、光照、水分胁迫、营养、CO2浓度、空气湿度）及植物的发育阶段、机械损伤和昆虫取食等对植物挥发性化合物合成与释放的影响机制。     

Modelling changes in VOC emission in response to climate change in the continental United States

The alteration of climate is driven not only by anthropogenic activities, but also by biosphere processes that change in conjunction with climate. Emission of volatile organic compounds (VOCs) from vegetation may be particularly sensitive to changes in climate and may play an important role in climate forcing through their influence on the atmospheric oxidative balance, greenhouse gas concentration, and the formation of aerosols. Using the VEMAP vegetation database and associated vegetation responses to climate change, this study examined the independent and combined effects of simulated changes in temperature, CO₂ concentration, and vegetation distribution on annual emissions of isoprene, monoterpenes, and other reactive VOCs (ORVOCs) from potential vegetation of the continental United States. Temperature effects were modelled according to the direct influence of temperature on enzymatic isoprene production and the vapour pressure of monoterpenes and ORVOCs. The effect of elevated CO₂ concentration was modelled according to increases in foliar biomass per unit of emitting surface area. The effects of vegetation distribution reflects simulated changes in species spatial distribution and areal coverage by 21 different vegetation classes. Simulated climate warming associated with a doubled atmospheric CO₂ concentration enhanced total modelled VOC emission by 81.8% (isoprene + 82.1%, monoterpenes + 81.6%, ORVOC + 81.1%), whereas a simulated doubled CO₂ alone enhanced total modelled VOC emission by only + 11.8% (isoprene + 13.7%, monoterpenes + 4.1%, ORVOC + 11.7%). A simulated redistribution of vegetation in response to altered temperatures and precipitation patterns caused total modelled VOC emission to decline by 10.4% (isoprene – 11.7%, monoterpenes – 18.6%, ORVOC 0.0%) driven by a decline in area covered by vegetation classes emitting VOCs at high rates. Thus, the positive effect of leaf-level adjustments to elevated CO₂ (i.e. increases in foliar biomass) is balanced by the negative effect of ecosystem-level adjustments to climate (i.e. decreases in areal coverage of species emitting VOC at high rates).     

The future of South East Asian rainforests in a changing landscape and climate

With a focus on the Danum Valley area of Sabah, Malaysian Borneo, this special issue has as its theme the future of tropical rainforests in a changing landscape and climate. The global environmental context to the issue is briefly given before the contents and rationale of the issue are summarized. Most of the papers are based on research carried out as part of the Royal Society South East Asia Rainforest Research Programme. The issue is divided into five sections: (i) the historical land-use and land management context; (ii) implications of land-use change for atmospheric chemistry and climate change; (iii) impacts of logging, forest fragmentation (particularly within an oil palm plantation landscape) and forest restoration on ecosystems and their functioning; (iv) the response and resilience of rainforest systems to climatic and land-use change; and (v) the scientific messages and policy implications arising from the research findings presented in the issue.     

Human-induced changes in US biogenic volatile organic compound emissions: evidence from long-term forest inventory data

Volatile organic compounds (VOCs) emitted by woody vegetation influence global climate forcing and the formation of tropospheric ozone. We use data from over 250 000 re‐surveyed forest plots in the eastern US to estimate emission rates for the two most important biogenic VOCs (isoprene and monoterpenes) in the 1980s and 1990s, and then compare these estimates to give a decadal change in emission rate. Over much of the region, particularly the southeast, we estimate that there were large changes in biogenic VOC emissions: half of the grid cells (1°× 1°) had decadal changes in emission rate outside the range ?2.3% to +16.8% for isoprene, and outside the range 0.2–17.1% for monoterpenes. For an average grid cell the estimated decadal change in heatwave biogenic VOC emissions (usually an increase) was three times greater than the decadal change in heatwave anthropogenic VOC emissions (usually a decrease, caused by legislation). Leaf‐area increases in forests, caused by anthropogenic disturbance, were the most important process increasing biogenic VOC emissions. However, in the southeast, which had the largest estimated changes, there were substantial effects of ecological succession (which decreased monoterpene emissions and had location‐specific effects on isoprene emissions), harvesting (which decreased monoterpene emissions and increased isoprene emissions) and plantation management (which increased isoprene emissions, and decreased monoterpene emissions in some states but increased monoterpene emissions in others). In any given region, changes in a very few tree species caused most of the changes in emissions: the rapid changes in the southeast were caused almost entirely by increases in sweetgum (Liquidambar styraciflua) and a few pine species. Therefore, in these regions, a more detailed ecological understanding of just a few species could greatly improve our understanding of the relationship between natural ecological processes, forest management, and biogenic VOC emissions.     

Fuelling the biodiversity crisis: species loss of ground-dwelling forest ants in oil palm plantations in Sabah,Malaysia (Borneo)

Oil palm plantations today cover large areas of former tropical lowland rain forest in Southeast Asia and are rapidly expanding on the island of Borneo. Study of the community of ground-dwelling ants in different plantations in Sabah, Malaysia, over 2 years using tuna baiting, revealed that the oil palm plantation ground ant community was severely reduced in species richness in comparison to the forest interior, regardless of age, undergrowth cover, or proximity to neighbouring forest. The results indicate that oil palm plantation habitats, now covering more than 15% of Sabah’s land area, can sustain only about 5% of the ground-dwelling ant species of the forest interior. Nine of the 23 ant species baited in the plantations were never recorded inside forest. All numerically dominant ants were non-forest species. The most common species was Anoplolepis gracilipes, an invasive species present at 70% of all bait sites and known to cause ‘ecological meltdowns’ in other situations. The low frequency and species number of forest ground ants indicates that oil palm plantations act as effective dispersal barriers leading to community isolation in rain forest remnants. The replacement of natural forests with oil palm plantations poses a serious threat to the conservation of biodiversity on Borneo if similar results are confirmed in other taxa.     

Parasitoid wasp communities on oil palm plantation: Effects of natural habitat existence are obscured by lepidopteran abundance

The existence of natural habitat around agroecosystem plays a pivotal role in maintaining the presence of natural enemies especially parasitoids. Natural habitat can fail to support biological pest control in agroecosystem due to a particular condition. The objective of this research was to investigate the effect of natural habitat existence on parasitoid wasp communities in oil palm plantation. The ecological research was conducted in the oil palm plantation located in Central Borneo, Indonesia. Twelve plots of oil palm plantations with different habitat characteristics were selected. Insects were sampled using canopy knockdown fogging with a pyrethroid insecticide. In total, 237 species and 2669 individuals of parasitoid wasps belong to 15 families were collected from all research areas. The results showed that species richness and abundance of parasitoid wasps in oil palm plantation was affected by lepidopteran abundance and not by the existence of natural habitat. However, the distance and area of natural habitat influenced the species composition of parasitoid wasps. We concluded that the existence of natural habitat still can maintain the parasitoid wasps in oil palm plantation. Thus, efforts on maintaining parasitoid wasp diversity for management of biological control in oil palm plantation need to pay attention by conserving the natural habitats.     

Amazonian biogenic volatile organic compounds under global change

Biogenic volatile organic compounds (BVOCs) play important roles at cellular, foliar, ecosystem and atmospheric levels. The Amazonian rainforest represents one of the major global sources of BVOCs, so its study is essential for understanding BVOC dynamics. It also provides insights into the role of such large and biodiverse forest ecosystem in regional and global atmospheric chemistry and climate. We review the current information on Amazonian BVOCs and identify future research priorities exploring biogenic emissions and drivers, ecological interactions, atmospheric impacts, depositional processes and modifications to BVOC dynamics due to changes in climate and land cover. A feedback loop between Amazonian BVOCs and the trends of climate and land‐use changes in Amazonia is then constructed. Satellite observations and model simulation time series demonstrate the validity of the proposed loop showing a combined effect of climate change and deforestation on BVOC emission in Amazonia. A decreasing trend of isoprene during the wet season, most likely due to forest biomass loss, and an increasing trend of the sesquiterpene to isoprene ratio during the dry season suggest increasing temperature stress‐induced emissions due to climate change.     

Barriers to dispersal of rain forest butterflies in tropical agricultural landscapes

Fragmentation of natural habitats can be detrimental for species if individuals fail to cross habitat boundaries to reach new locations, thereby reducing functional connectivity. Connectivity is crucial for species shifting their ranges under climate change, making it important to understand factors that might prevent movement through human‐modified landscapes. In tropical regions, rain forests are being fragmented by agricultural expansion, potentially isolating populations of highly diverse forest‐dependent species. The likelihood of crossing habitat boundaries is an important determinant of species dispersal through fragmented landscapes, and so we examined movement across rain forest‐oil palm plantation boundaries on Borneo by using relatively mobile nymphalid butterflies as our model study taxon. We marked 1666 individuals from 65 species, and 19 percent (100/527) of recaptured individuals crossed the boundary. Boundary crossing was relatively frequent in some species, and net movement of individuals was from forest into plantation. However, boundary crossing from forest into plantation was detected in less than 50 percent (12/28) of recaptured species and was dominated by small‐sized butterfly species whose larval host plants occurred within plantations. Thus, while oil palm plantations may be relatively permeable to some species, they may act as barriers to the movement of forest‐dependent species (i.e., species that require rain forest habitat to breed), highlighting the importance of maintaining forest connectivity for conserving rain forest species.     

浙江省毛竹异戊二烯排放规律及其影响

主要对中国南方广泛分布的毛竹进行了异戊二烯排放的观测和模拟研究 ,以浙江省为例进行了毛竹异戊二烯排放量的估算 ,并将太湖流域的竹子分布与 O3浓度观测结果进行了比较 ,以期探索毛竹对区域对流层大气化学环境变化的贡献。观测结果表明毛竹是异戊二烯排放潜力较大的植被 ,其值为 1 1 6± 2 3 .4ug.g- 1.h- 1,浙江省 1 996年毛竹的异戊二烯排放量为 0 .75 9Mkg,排放在夏季达到峰值 ,其中 6、7、8三个月的排放量占全年总排放量的 5 8.9%。通过将 O3观测结果与竹林分布比较分析推断 ,农村地区 (临安 )出现的高 O3浓度可能与大面积竹林的异戊二烯排放有关。如果这种假设能够得到证明 ,将为一些农村地区大气高臭氧浓度的形成提供一种理论解释     

Rainforest conversion to smallholder cash crops leads to varying declines of beetles (Coleoptera) on Sumatra

Southeast Asian arthropod biodiversity is in rapid decline, but the variability of responses within taxa has received little attention. Using canopy fogging, we collected ~50,000 beetles (Coleoptera) in (1) lowland rainforest, (2) jungle rubber (rubber agroforest), and smallholder monoculture plantations of (3) rubber and (4) oil palm in Sumatra, across two landscapes and seasons. On average, beetle abundance was more than 50%, and biomass over 75%, lower in rubber and oil palm plantations than in rainforest and jungle rubber. This pattern was influenced by landscape and season. Abundance and biomass declines were similar in Chrysomelidae, Elateridae, and Staphylinidae, but differed in Curculionidae, which were most abundant in oil palm due to the introduced oil palm pollinator Elaeidobius kamerunicus. Across beetle families, species richness in monocultures was reduced by at least 70% compared to rainforest, with beetle richness in jungle rubber being similar to rainforest. Community composition in oil palm plantations differed markedly from the other land-use systems for Chrysomelidae and Curculionidae, but less for Elateridae and Staphylinidae. Turnover contributed more to overall beta diversity than nestedness for all families and land-use systems. Likely undersampling of the beetle community in rainforest suggests that declines of beetle density and diversity are much more severe than reported here, especially for beetle families with many concealed species, such as Staphylinidae. This study provides first evidence that negative responses of beetles to tropical land-use change vary among families, and is the first report of its kind from heavily understudied Sumatra.     

Circadian control of isoprene emissions from oil palm (Elaeis guineensis)

The emission of isoprene from the biosphere to the atmosphere has a profound effect on the Earth's atmospheric system. Until now, it has been assumed that the primary short-term controls on isoprene emission are photosynthetically active radiation and temperature. Here we show that isoprene emissions from a tropical tree (oil palm, Elaeis guineensis) are under strong circadian control, and that the circadian clock is potentially able to gate light-induced isoprene emissions. These rhythms are robustly temperature compensated with isoprene emissions still under circadian control at 38 degrees C. This is well beyond the acknowledged temperature range of all previously described circadian phenomena in plants. Furthermore, rhythmic expression of LHY/CCA1, a genetic component of the central clock in Arabidopsis thaliana, is still maintained at these elevated temperatures in oil palm. Maintenance of the CCA1/LHY-TOC1 molecular oscillator at these temperatures in oil palm allows for the possibility that this system is involved in the control of isoprene emission rhythms. This study contradicts the accepted theory that isoprene emissions are primarily light-induced.     

Within‐plant isoprene oxidation confirmed by direct emissions of oxidation products methyl vinyl ketone and methacrolein

Isoprene is emitted from many terrestrial plants at high rates, accounting for an estimated 1/3 of annual global volatile organic compound emissions from all anthropogenic and biogenic sources combined. Through rapid photooxidation reactions in the atmosphere, isoprene is converted to a variety of oxidized hydrocarbons, providing higher order reactants for the production of organic nitrates and tropospheric ozone, reducing the availability of oxidants for the breakdown of radiatively active trace gases such as methane, and potentially producing hygroscopic particles that act as effective cloud condensation nuclei. However, the functional basis for plant production of isoprene remains elusive. It has been hypothesized that in the cell isoprene mitigates oxidative damage during the stress‐induced accumulation of reactive oxygen species (ROS), but the products of isoprene‐ROS reactions in plants have not been detected. Using pyruvate‐2‐¹³C leaf and branch feeding and individual branch and whole mesocosm flux studies, we present evidence that isoprene (i) is oxidized to methyl vinyl ketone and methacrolein (i_ox) in leaves and that i_ox/i emission ratios increase with temperature, possibly due to an increase in ROS production under high temperature and light stress. In a primary rainforest in Amazonia, we inferred significant in plant isoprene oxidation (despite the strong masking effect of simultaneous atmospheric oxidation), from its influence on the vertical distribution of i_ox uptake fluxes, which were shifted to low isoprene emitting regions of the canopy. These observations suggest that carbon investment in isoprene production is larger than that inferred from emissions alone and that models of tropospheric chemistry and biota–chemistry–climate interactions should incorporate isoprene oxidation within both the biosphere and the atmosphere with potential implications for better understanding both the oxidizing power of the troposphere and forest response to climate change.     

Effects of Logging and Oil Palm Expansion on Stream Frog Communities on Borneo,Southeast Asia

The conversion of tropical rain forests to oil palm plantations is a major threat to Southeast Asia's rich biodiversity. Fostering forest species communities in secondary forests, agroforestry systems, and plantations is therefore increasingly becoming a conservation focus. This study uses standardized transect‐based sampling to compare species richness, density and community composition of stream anuran assemblages among primary forests, repeatedly logged forests and oil palm plantations in northern Borneo. In primary forest streams, we recorded an average of 19 frog species, compared to 15 species in logged forests and 11 species in oil palm plantation streams. However, the high percentage of canopy cover above the plantation streams mitigated this loss to some extent. This study corroborates numerous studies that oil palm plantations have mainly negative effects on the region's biodiversity. However, our results also demonstrate the high conservation value of logged forests for Bornean stream‐dependent anurans. We conclude that palm plantations have a largely unused potential to promote regional anuran biodiversity.     

Heterogeneity of tree diversity and carbon stocks in Amazonian oil palm landscapes

ABSTRACT

Background: Quantitative effects of large-scale oil palm expansion in the Neotropics on biodiversity and carbon stocks are still poorly documented.

Aims: We evaluated differences in tree species composition and richness, and above-ground carbon stocks among dominant land cover types in Pará state, Brazil.

Methods: We quantified tree species composition and richness and above-ground carbon stock in stands in remnant primary rain forest, young secondary forest, oil palm plantation and pastures.

Results: We sampled 5,696 trees with a DBH ≥ 2 cm, of 413 species in 68 families, of which 381 species were recorded in primary forest fragments. We found significant differences in species richness and carbon stock among the four land cover classes. Carbon stocks in remnant primary forest were typically over 190 Mg ha^?1, while those in other land cover types were typically less than 60 Mg ha^?1.

Conclusion: Oil palm plantations have a species-poor tree community given active management; old plantations have a standing carbon stock which is comparable to that of secondary forest and far greater than that of pastures. Private forest reserves within oil palm company holdings play an important role in preserving primary forest tree diversity in human-modified landscapes in Amazonia.     

Conclusion: applying South East Asia Rainforest Research Programme science to land-use management policy and practice in a changing landscape and climate

The context and challenges relating to the remaining tropical rainforest are briefly reviewed and the roles which science can play in addressing questions are outlined. Key messages which articles in the special issue, mainly based on projects of the Royal Society South East Asia Rainforest Research Programme (SEARRP), have raised of relevance to policies on land use, land management and REDD+ are then considered. Results from the atmospheric science and hydrology papers, and some of the ecological ones, demonstrate the very high ecosystem service values of rainforest (compared with oil palm) in maintaining high biodiversity, good local air quality, reducing greenhouse emissions, and reducing landslide, flooding and sedimentation consequences of climate change-and hence provide science to underpin the protection of remaining forest, even if degraded and fragmented. Another group of articles test ways of restoring forest quality (in terms of biodiversity and carbon value) or maintaining as high biodiversity and ecological functioning levels as possible via intelligent design of forest zones and fragments within oil palm landscapes. Finally, factors that have helped to enhance the policy relevance of SEARRP projects and dissemination of their results to decision-makers are outlined.     

Impacts of logging and conversion of rainforest to oil palm on the functional diversity of birds in Sundaland

Commercial selective logging and the conversion of primary and degraded forests to agriculture are the biggest threats to tropical biodiversity. Our understanding of the impacts of these disturbances and the resulting local extinctions on the functional roles performed by the remaining species is limited. We address this issue by examining functional diversity (FD), which quantifies a range of traits that affect a species' ecological role in a community as a single continuous metric. We calculated FD for birds across a gradient of disturbance from primary forest through intensively logged forest to oil palm plantations on previously forested land in Borneo, Southeast Asia, a hotspot of imperilled biodiversity. Logged rainforest retained similar levels of FD to unlogged rainforest, even after two logging rotations, but the conversion of logged forest to oil palm resulted in dramatic reductions in FD. The few remaining species in oil palm filled a disproportionately wide range of functional roles but showed very little clustering in terms of functional traits, suggesting that any further extinctions from oil palm would reduce FD even further. Determining the extent to which the changes we recorded were due to under‐utilization of resources within oil palm or a reduction in the resources present is an important next step. Nonetheless our study improves our understanding of the stability and resilience of functional diversity in these ecosystems and of the implications of land‐use changes for ecosystem functioning.     

The Impact of Selective-Logging and Forest Clearance for Oil Palm on Fungal Communities in Borneo

Tropical forests are being rapidly altered by logging, and cleared for agriculture. Understanding the effects of these land use changes on soil fungi, which play vital roles in the soil ecosystem functioning and services, is a major conservation frontier. Using 454-pyrosequencing of the ITS1 region of extracted soil DNA, we compared communities of soil fungi between unlogged, once-logged, and twice-logged rainforest, and areas cleared for oil palm, in Sabah, Malaysia. Overall fungal community composition differed significantly between forest and oil palm plantation. The OTU richness and Chao 1 were higher in forest, compared to oil palm plantation. As a proportion of total reads, Basidiomycota were more abundant in forest soil, compared to oil palm plantation soil. The turnover of fungal OTUs across space, true β-diversity, was also higher in forest than oil palm plantation. Ectomycorrhizal (EcM) fungal abundance was significantly different between land uses, with highest relative abundance (out of total fungal reads) observed in unlogged forest soil, lower abundance in logged forest, and lowest in oil palm. In their entirety, these results indicate a pervasive effect of conversion to oil palm on fungal community structure. Such wholesale changes in fungal communities might impact the long-term sustainability of oil palm agriculture. Logging also has more subtle long term effects, on relative abundance of EcM fungi, which might affect tree recruitment and nutrient cycling. However, in general the logged forest retains most of the diversity and community composition of unlogged forest.     

Survival strategies of a frugivore,the sun bear,in a forest-oil palm landscape

As large areas of forest are lost throughout the tropics, prime habitat of many species decline and become fragmented. The island of Borneo is a prime example, with accelerated clearing of forests primarily for oil palm expansion. Borneo is recognized as an important stronghold for the conservation of the sun bear (Helarctos malayanus), but it is unclear how habitat reduction and fragmentation is affecting this frugivore. We used camera traps and sign surveys to understand patterns of sun bear habitat use in a matrix of fragmented forests and extensive oil palm development, which has existed as such for?>?15 years: the Lower Kinabatangan floodplain in Sabah, Malaysian Borneo. Within these small forest fragments, squeezed between a major river and oil palm plantations, bears exhibited selection for areas farther from human activity (plantations, river boat traffic, and buildings), and were rarely active during the day, demonstrating both spatial and temporal avoidance of potential human-related threats. They selected large trees to feed and rest, and also exploited adjacent plantations to feed on oil palm fruits. We conclude that even relatively small forest fragments (~?2000 ha) within large agricultural landscapes can be important for sun bears. Our research highlights the remarkable adaptations this species has employed to persist in a drastically modified landscape.