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Aims Non-structural carbohydrates (NSCs) are plant storage compounds used for metabolism, transport, osmoregulation and regrowth following the loss of plant tissue. Even in conditions suitable for optimal growth, plants continue to store NSCs. This storage may be due to passive accumulation from sink-inhibited growth or active reserves that come at the expense of growth. The former pathway implies that NSCs may be a by-product of sink limitation, while the latter suggests a functional role of NSCs for use during poor conditions. Methods Using 13C pulse labelling, we traced the source of soluble sugars in stem and root organs during drought and everwet conditions for seedlings of two tropical tree species that differ in drought tolerance to estimate the relative allocation of NSCs stored prior to drought versus NSCs assimilated during drought. We monitored growth, stomatal conductance, stem water potential and NSC storage to assess a broad carbon response to drought. Important Findings We found that the drought-sensitive species had reduced growth, conserved NSC concentrations in leaf, stem and root organs and had a larger proportion of soluble sugars in stem and root organs that originated from pre-drought storage relative to seedlings in control conditions. In contrast, the drought-tolerant species maintained growth and stem and root NSC concentrations but had reduced leaf NSCs concentrations with a larger proportion of stem and root soluble sugars originated from freshly assimilated photosynthates relative to control seedlings. These results suggest the drought-sensitive species passively accumulated NSCs during water deficit due to growth inhibition, while the drought-tolerant species actively responded to water deficit by allocating NSCs to stem and root organs. These strategies seem correlated with baseline maximum growth rates, which supports previous research suggesting a trade-off between growth and drought tolerance while providing new evidence for the importance of plasticity in NSC allocation during drought.

Patterns of co-occurrence among species can help reveal the structure and assembly of ecological communities. However, studies have been limited by measuring co-occurrence in either space or time but not both simultaneously. This is especially problematic in systems such as masting forests where resources are highly variable, meaning that spatial use and co-occurrence patterns can change on fine spatiotemporal scales. We develop an analytical framework for assessing species co-occurrence at fine spatial and temporal scales simultaneously and apply these models to a camera trapping dataset from Borneo. We sought to determine how substantial variation in food availability across space and time affects co-occurrence among terrestrial vertebrates. We detect many significant, mostly positive, co-occurrence patterns among species, but almost entirely in unlogged forest and during dipterocarp mast years. The most strongly co-occurring pair of species, bearded pig (Sus barbatus) and sambar (Rusa unicolor), only positively co-occur in areas and years when fruit is locally abundant. Species occurrences in logged forest and non-mast years are mostly random with respect to other species. This suggests that frugivore-granivore species positively co-occur when resources are plentiful (i.e., large trees are present and fruiting), likely because they use the same resources; these patterns disappear when food availability is lower. Our approach demonstrates the utility of measuring co-occurrence in space and time together and highlights the importance of resource abundance for driving the co-occurrence structure of communities. Furthermore, our method could be broadly applied to other systems to assess fine-scale spatiotemporal patterns across a range of taxa.

Overhunting reduces important plant-animal interactions such as vertebrate seed dispersal and seed predation, thereby altering plant regeneration and even above-ground biomass. It remains unclear, however, if non-hunted species can compensate for lost vertebrates in defaunated ecosystems. We use a nested exclusion experiment to isolate the effects of different seed enemies in a Bornean rainforest. In four of five tree species, vertebrates kill many seeds (13–66%). Nonetheless, when large mammals are excluded, seed mortality from insects and fungi fully compensates for the lost vertebrate predation, such that defaunation has no effect on seedling establishment. The switch from seed predation by generalist vertebrates to specialist insects and fungi in defaunated systems may alter Janzen–Connell effects and density-dependence in plants. Previous work using simulation models to explore how lost seed dispersal will affect tree species composition and carbon storage may require reevaluation in the context of functional redundancy within complex species interactions networks., Ecologists predict that loss of large vertebrates will alter tropical plant communities. Here, the authors report a field experiment on seed mortality and seedling establishment in Borneo, in which experimental defaunation of large seed consumers was functionally compensated by insects and fungi.

Occasional periods of drought are typical of most tropical forests, but climate change is increasing drought frequency and intensity in many areas across the globe, threatening the structure and function of these ecosystems. The effects of intermittent drought on tropical tree communities remain poorly understood and the potential impacts of intensified drought under future climatic conditions are even less well known. The response of forests to altered precipitation will be determined by the tolerances of different species to reduced water availability and the interactions among plants that alleviate or exacerbate the effects of drought. Here, we report the response of experimental monocultures and mixtures of tropical trees to simulated drought, which reveals a fundamental shift in the nature of interactions among species. Weaker competition for water in diverse communities allowed seedlings to maintain growth under drought while more intense competition among conspecifics inhibited growth under the same conditions. These results show that reduced competition for water among species in mixtures mediates community resistance to drought. The delayed onset of competition for water among species in more diverse neighbourhoods during drought has potential implications for the coexistence of species in tropical forests and the resilience of these systems to climate change. Reduced competition for water among species in mixed tropical plant communities mediates community resistance to drought: weaker competition permits growth maintenance in drought, whereas stronger competition inhibits it.

Precipitation patterns are changing across the globe causing more severe and frequent drought for many forest ecosystems. Although research has focused on the resistance of tree populations and communities to these novel precipitation regimes, resilience of forests is also contingent on recovery following drought, which remains poorly understood, especially in aseasonal tropical forests. We used rainfall exclusion shelters to manipulate the interannual frequency of drought for diverse seedling communities in a tropical forest and assessed resistance, recovery and resilience of seedling growth and mortality relative to everwet conditions. We found seedlings exposed to recurrent periods of drought altered their growth rates throughout the year relative to seedlings in everwet conditions. During drought periods, seedlings grew slower than seedlings in everwet conditions (i.e., resistance phase) while compensating with faster growth after drought (i.e., recovery phase). However, the response to frequent drought was species dependent as some species grew significantly slower with frequent drought relative to everwet conditions while others grew faster with frequent drought due to overcompensating growth during the recovery phase. In contrast, mortality was unrelated to rainfall conditions and instead correlated with differences in light. Intra-annual plasticity of growth and increased annual growth of some species led to an overall maintenance of growth rates of tropical seedling communities in response to more frequent drought. These results suggest these communities can potentially adapt to predicted climate change scenarios and that plasticity in the growth of species, and not solely changes in mortality rates among species, may contribute to shifts in community composition under drought.

Aim Financial incentives to manage forests sustainably, such as certification or carbon storage payments, are assumed to have co-benefits for biodiversity conservation. This claim remains little studied for rain forest mammals, which are particularly threatened, but challenging to survey. Location Sabah, Malaysia, Borneo. Methods We used photographic data from three commercial forest reserves to show how community occupancy modelling can be used to quantify mammalian diversity conservation co-benefits of forest certification. These reserves had different management histories, and one was certified by the Forest Stewardship Council. Results Many threatened species occupied larger areas in the certified reserve. Species richness, estimated per 200 × 200-m grid cell throughout all reserves, was higher in the certified site, particularly for threatened species. The certified reserve held the highest aboveground biomass. Within reserves, aboveground biomass was not strongly correlated with patterns of mammal richness (Spearman's rho from 0.03 to 0.32); discrepancies were strongest along reserve borders. Main conclusions Our approach provides a flexible and standardized tool to assess biodiversity and identify winners of sustainable forestry. Inferring patterns of species richness from camera-trapping carries potential for the objective designation of high conservation value forest. Correlating species richness with aboveground biomass further allows evaluating the biodiversity co-benefits of carbon protection. These advantages make the present approach an ideal tool to overcome the difficulties to rigorously quantify biodiversity co-benefits of forest certification and carbon storage payments.

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.

Conversion of tropical forests to oil palm plantations in Malaysia and Indonesia has resulted in large-scale environmental degradation, loss of biodiversity and significant carbon emissions. For both countries to participate in the United Nation’s REDD (Reduced Emission from Deforestation and Degradation) mechanism, assessment of forest carbon stocks, including the estimated loss in carbon from conversion to plantation, is needed. In this study, we use a combination of field and remote sensing data to quantify both the magnitude and the geographical distribution of carbon stock in forests and timber plantations, in Sabah, Malaysia, which has been the site of significant expansion of oil palm cultivation over the last two decades. Forest structure data from 129 ha of research and inventory plots were used at different spatial scales to discriminate forest biomass across degradation levels. Field data was integrated with ALOS PALSAR (Advanced Land-Observing Satellite Phased Array L-band Synthetic Aperture Radar) imagery to both discriminate oil palm plantation from forest stands, with an accuracy of 97.0% (κ = 0.64) and predict AGB using regression analysis of HV-polarized PALSAR data (R² = 0.63, p < .001). Direct estimation of AGB from simple regression models was sensitive to both environmental conditions and forest structure. Precipitation effect on the backscatter data changed the HV prediction of AGB significantly (R² = 0.21, p < .001), and scattering from large leaves of mature palm trees significantly impeded the use of a single HV-based model for predicting AGB in palm oil plantations. Multi-temporal SAR data and algorithms based on forest types are suggested to improve the ability of a sensor similar to ALOS PALSAR for accurately mapping and monitoring forest biomass, now that the ALOS PALSAR sensor is no longer operational.

In a shadehouse experiment we tested the effects of light nutrients and ectomycorrhizal fungi (EMF) on the growth of Vatica albiramis van Slooten (Dipterocarpaceae) seedlings. We hypothesised that it is more advantageous for plants to form connections with EMF and to trade carbon for nutrients with EMF under high light than low light. The relationship between seedling growth and the proportion of ectomycorrhizal root tips was expected as positive in high light and as negative in low light. Light conditions simulated the forest understory (low; 3 full sunlight) a small gap (medium; 11) and a large gap (high; 33) and a fully factorial combination of nutrients (F-/+) and ectomycorrhizal colonization (EMF-/+) treatments were applied within light conditions. The application of EMF and nutrients did significantly alter seedling growth across the range of forest floor light conditions however the key hypothesis was rejected as seedling growth under low light was not affected by increased EMF colonization of root tips (light:EMF colonization ?2?=?2.97 p? =?0.23). In addition the lack of difference in morphotype abundance across light conditions indicated that light changes may not favour the association to specific EMF in seedlings of this particular dipterocarp species. Our results suggest that antagonistic (non beneficial to the plant) effects due to ectomycorrhizal colonization under a light constrained environment may not affect seedling growth of Vatica albiramis. Electronic supplementary material: The online version of this article (doi:10.1007/s11104 010 0555 3) contains supplementary material which is available to authorized users.

BACKGROUND: The Janzen-Connell hypothesis proposes that seed and seedling enemies play a major role in maintaining high levels of tree diversity in tropical forests. However, human disturbance may alter guilds of seed predators including their body size distribution. These changes have the potential to affect seedling survival in logged forest and may alter forest composition and diversity. METHODOLOGY/PRINCIPAL FINDINGS: We manipulated seed density in plots beneath con- and heterospecific adult trees within a logged forest and excluded vertebrate predators of different body sizes using cages. We show that small and large-bodied predators differed in their effect on con- and heterospecific seedling mortality. In combination small and large-bodied predators dramatically decreased both con- and heterospecific seedling survival. In contrast, when larger-bodied predators were excluded small-bodied predators reduced conspecific seed survival leaving seeds coming from the distant tree of a different species. CONCLUSIONS/SIGNIFICANCE: Our results suggest that seed survival is affected differently by vertebrate predators according to their body size. Therefore, changes in the body size structure of the seed predator community in logged forests may change patterns of seed mortality and potentially affect recruitment and community composition

To evaluate the effectiveness of sustainable forest management (SFM) for wildlife conservation, we investigated the abundances of medium to large ground-dwelling vertebrates in a forest management unit in Borneo by camera trapping. The forest management unit (FMU), Deramakot Forest Reserve (55,083 ha), has applied SFM for the past 15 years. We established 15 plots in preharvested areas and five plots in postharvested areas over the FMU. Plots in the postharvested areas had been subject to reduced-impact logging from 2 to 13 years ago. We obtained photos of ground-dwelling vertebrates with infrared sensor cameras set at 12 random points in each plot. Based on the numbers of photos taken over 770 camera days in each plot, we calculated the mean trapping rate (MTR) of each species for each plot. Over the 20 plots, we obtained 5444 photos of 39 medium-to-large vertebrates (i.e., mammals, birds, and monitor lizards); these included many elusive and endangered species. Among the 39 species, no species showed a significant difference in MTR between the pre- and postharvested areas. Furthermore, species composition was not significantly different between the pre- and postharvested areas. Our results support the idea that implementation of SFM can be an effective investment in wildlife conservation in tropical rainforests.

Borneo has experienced a rapid decline in the extent of forest cover, which has reduced the amount of habitat available for many plant and animal species. The precise impact of habitat loss on the conservation status of dipterocarp trees is uncertain. We use three contrasting techniques, the extent of occurrence, area of occupancy and ecological niche models derived using maxent, in conjunction with a current land-use map of Sabah, to derive estimates of habitat loss and infer a regional IUCN Red List conservation status for 33 Sabah dipterocarp species. Estimates of habitat loss differed significantly according to the methods employed and between species on different habitat types. Proportion of habitat loss determined from the ecological niche models varied from 21 percent for Shorea micans to 99.5 percent for Dipterocarpus lamellatus. Thirty-two of the 33 dipterocarp species analyzed in this study would have their Sabah populations classified as Threatened (equal to a habitat loss of > 30%) under the A2 IUCN Red List criterion. Dipterocarps that occur in lowland forests have experienced greater habitat loss than upland/lower montane or ultramafic species. In addition, species with the lowest predicted area within their historic distributions had the highest proportion of habitat lost, which provides a rationale for targeting conservation effort on the species with narrow distributions. We recommend the ecological niche modeling approach as a rapid assessment tool for reconstructing species’ historic distributions during conservation assessments of tropical trees.

Stable carbon isotope ( δ 13 C) series were developed from analysis of sequential radial wood increments from AD 1850 to AD 2009 for four mature primary rainforest trees from the Danum and Imbak areas of Sabah, Malaysia. The aseasonal equatorial climate meant that conventional dendrochronology was not possible as the tree species investigated do not exhibit clear annual rings or dateable growth bands. Chronology was established using radiocarbon dating to model age–growth relationships and date the carbon isotopic series from which the intrinsic water-use efficiency (IWUE) was calculated. The two Eusideroxylon zwageri trees from Imbak yielded ages of their pith/central wood (±1 sigma) of 670 ± 40 and 759 ± 40 years old; the less dense Shorea johorensis and Shorea superba trees at Danum yielded ages of 240 ± 40 and 330 ± 40 years, respectively. All trees studied exhibit an increase in the IWUE since AD 1960. This reflects, in part, a response of the forest to increasing atmospheric carbon dioxide concentration. Unlike studies of some northern European trees, no clear plateau in this response was observed. A change in the IWUE implies an associated modification of the local carbon and/or hydrological cycles. To resolve these uncertainties, a shift in emphasis away from high-resolution studies towards long, well-replicated time series is proposed to develop the environmental data essential for model evaluation. Identification of old (greater than 700 years) ringless trees demonstrates their potential in assessing the impacts of climatic and atmospheric change. It also shows the scientific and applied value of a conservation policy that ensures the survival of primary forest containing particularly old trees (as in Imbak Canyon and Danum).

The carbon storage and conservation value of old-growth tropical forests is clear, but the value of logged forest is less certain. Here we analyse >100,000 observations of individuals from 11 taxonomic groups and >2,500 species, covering up to 19 years of post-logging regeneration, and quantify the impacts of logging on carbon storage and biodiversity within lowland dipterocarp forests of Sabah, Borneo. We estimate that forests lost ca. 53% of above-ground biomass as a result of logging but despite this high level of degradation, logged forest retained considerable conservation value: floral species richness was higher in logged forest than in primary forest and whilst faunal species richness was typically lower in logged forest, in most cases the difference between habitats was no greater than ca. 10%. Moreover, in most studies >90% of species recorded in primary forest were also present in logged forest, including species of conservation concern. During recovery, logged forest accumulated carbon at five times the rate of natural forest (1.4 and 0.28 Mg C ha−1 year−1, respectively). We conclude that allowing the continued regeneration of extensive areas of Borneo’s forest that have already been logged, and are at risk of conversion to other land uses, would provide a significant carbon store that is likely to increase over time. Protecting intact forest is critical for biodiversity conservation and climate change mitigation, but the contribution of logged forest to these twin goals should not be overlooked.

In this paper previously unpublished field data from 25 ha of permanent sampling plots (PSPs) in Sabah, Malaysia, in four different forest reserves are analysed for mortality rates and basal area development. Field data of an observation length of 9 to 20 years were available. These data then form the basis of several benchmark tests for the evaluation of the individual-oriented tropical rain forest growth model formind . A new version of the formind is presented. The model in its version formind1.1 includes enhanced submodels for mortality and tree growth. The model evaluation is focused on the model components for tree growth, competition and mortality. Data for tree recruitment were not available. Results show a good agreement between simulation and field data for the main output variables basal area and stem number indicating a reasonable behaviour of the model components we focused on. Furthermore the results show that differences in site conditions influence tree growth and mortality. Site characteristics should be included in the model in the future.

Deforestation in the tropics is an important source of carbon C release to the atmosphere. To provide a sound scientific base for efforts taken to reduce emissions from deforestation and degradation (REDD+) good estimates of C stocks and fluxes are important. We present components of the C balance for selectively logged lowland tropical dipterocarp rainforest in the Malua Forest Reserve of Sabah, Malaysian Borneo. Total organic C in this area was 167.9 Mg C ha⁻¹±3.8 (SD), including: Total aboveground (TAGC: 55%; 91.9 Mg C ha⁻¹±2.9 SEM) and belowground carbon in trees (TBGC: 10%; 16.5 Mg C ha⁻¹±0.5 SEM), deadwood (8%; 13.2 Mg C ha⁻¹±3.5 SEM) and soil organic matter (SOM: 24%; 39.6 Mg C ha⁻¹±0.9 SEM), understory vegetation (3%; 5.1 Mg C ha⁻¹±1.7 SEM), standing litter (

Tropical forests sequester a large amount of carbon in the vegetation and soils (approximately 25 % of the carbon in the terrestrial ecosystems) (Bonan 2008). The conversion of tropical forests through forestry and land-use change results in an enormous emission of carbon to the atmosphere. In the past decade, deforestation and forest degradation in Southeast Asia and elsewhere in the tropics accounted for approximately 20 % of global anthropogenic carbon emissions (Gullison et al. 2007). The important roles of natural tropical forests in carbon storage and climate change mitigation are increasingly recognized.

Comparisons between the consequences of improved management and those of conventional logging in terms of carbon stock and biodiversity conservation were conducted in the Deramakot Forest Reserve versus the Tangkulap Forest Reserve, where lowland dipterocarp forest is the natural vegetation (see also Imai et al. 2009). In this chapter, we summarize the co-benefits of improved forest management for biodiversity conservation and carbon sequestration. We evaluate the co-benefits in terms of additionality, which is the net increment effect of the improved forest management added over 13 years from 1989 (when logging was suspended in Deramakot) until 2002 (when our analyses started) in comparison to continued high-impact conventional logging.

The Deramakot Forest Reserve and the Tangklap Forest Reserve, where the ecological studies described in this book were conducted, are located in the interior of the state of Sabah, Malaysia (north Borneo) (Fig. 1.1). In Sabah, forested lands, including both natural and planted forests, are found mainly within areas designated by law as Forest Reserve, State Park, or Wildlife Sanctuary. Together these three classes of forest lands account for 53 % (3.87 million ha) of Sabah’s landmass, and are collectively referred to as the Permanent Forest Estate (PFE). Little native forest is currently left outside the boundaries of the PFE. Forest Reserve is managed under the Sabah Forestry Department and is designated by law primarily to produce timber from natural forests. Despite declining timber output, forestry remains a dominant land use in Sabah.

We studied the effects of forest composition and structure, and related biotic and abiotic factors on soil respiration rates in a tropical logged forest in Malaysian Borneo. Forest stands were classified into gap, pioneer, non-pioneer and mixed (pioneer, non-pioneer and unclassified trees) based on the species composition of trees >10 cm diameter breast height. Soil respiration rates did not differ significantly between non-gap sites (1290 ± 210 mg CO2 m-2 h-1) but were double those in gap sites (640 ± 130 mg CO2 m-2 h-1). Post hoc analyses found that an increase in soil temperature and a decrease in litterfall and fine root biomass explained 72% of the difference between gap and non-gap sites. The significant decrease of soil respiration rates in gaps, irrespective of day or night time, suggests that autotrophic respiration may be an important contributor to total soil respiration in logged forests. We conclude that biosphere-atmosphere carbon exchange models in tropical systems should incorporate gap frequency and that future research in tropical forest should emphasize the contribution of autotrophic respiration to total soil respiration. © 2009 Elsevier B.V. All rights reserved.

BackgroundSustainable forest management (SFM), which has been recently introduced to tropical natural production forests, is beneficial in maintaining timber resources, but information about the co-benefits for biodiversity conservation and carbon sequestration is currently lacking.Methodology/principal findingsWe estimated the diversity of medium to large-bodied forest-dwelling vertebrates using a heat-sensor camera trapping system and the amount of above-ground, fine-roots, and soil organic carbon by a combination of ground surveys and aerial-imagery interpretations. This research was undertaken both in SFM applied as well as conventionally logged production forests in Sabah, Malaysian Borneo. Our carbon estimation revealed that the application of SFM resulted in a net gain of 54 Mg C ha(-1) on a landscape scale. Overall vertebrate diversity was greater in the SFM applied forest than in the conventionally logged forest. Specifically, several vertebrate species (6 out of recorded 36 species) showed higher frequency in the SFM applied forest than in the conventionally logged forest.Conclusions/significanceThe application of SFM to degraded natural production forests could result in greater diversity and abundance of vertebrate species as well as increasing carbon storage in the tropical rain forest ecosystems.

Selective logging of tropical forests imposes spatial pattern on the landscape by creating a mosaic of patches affected by different intensities of disturbance. To understand the ecological impacts of selective logging it is therefore necessary to explore how patterns of tree species composition are affected by this patchy disturbance. This study examines the impacts of selective logging on species composition and spatial patterns of vegetation structure and tree diversity in Sabah, Borneo. We compare tree diversity between logged and unlogged forest at three scales: species richness within plots, species turnover among plots, and total species richness and composition of plots combined. Logging had no effect on tree diversity measured at the smallest scale. Logged forest had a greater rate of species turnover with distance, so at a large spatial scale it supported more tree species than the relatively homogeneous unlogged area. Tree species composition also differed significantly between the two types of forest, with more small dipterocarps and large pioneers in logged forest, and more large dipterocarps in unlogged forest. Our results emphasize the importance of sampling at a sufficiently large scale to represent patterns of biodiversity within tropical forest landscapes. Large areas of production forest in SE Asia are threatened with conversion to commercial crops; our findings show that selectively logged forest can retain considerable conservation value.