Thresholds for adding degraded tropical forest to the conservation estate.

Logged and disturbed forests are often viewed as degraded and depauperate environments compared with primary forest. However, they are dynamic ecosystems1 that provide refugia for large amounts of biodiversity2,3, so we cannot afford to underestimate their conservation value4. Here we present empirically defined thresholds for categorizing the conservation value of logged forests, using one of the most comprehensive assessments of taxon responses to habitat degradation in any tropical forest environment. We analysed the impact of logging intensity on the individual occurrence patterns of 1,681 taxa belonging to 86 taxonomic orders and 126 functional groups in Sabah, Malaysia. Our results demonstrate the existence of two conservation-relevant thresholds. First, lightly logged forests (<29% biomass removal) retain high conservation value and a largely intact functional composition, and are therefore likely to recover their pre-logging values if allowed to undergo natural regeneration. Second, the most extreme impacts occur in heavily degraded forests with more than two-thirds (>68%) of their biomass removed, and these are likely to require more expensive measures to recover their biodiversity value. Overall, our data confirm that primary forests are irreplaceable5, but they also reinforce the message that logged forests retain considerable conservation value that should not be overlooked.

Tropical forests post-logging are a persistent net carbon source to the atmosphere.

Logged and structurally degraded tropical forests are fast becoming one of the most prevalent land-use types throughout the tropics and are routinely assumed to be a net carbon sink because they experience rapid rates of tree regrowth. Yet this assumption is based on forest biomass inventories that record carbon stock recovery but fail to account for the simultaneous losses of carbon from soil and necromass. Here, we used forest plots and an eddy covariance tower to quantify and partition net ecosystem CO2 exchange in Malaysian Borneo, a region that is a hot spot for deforestation and forest degradation. Our data represent the complete carbon budget for tropical forests measured throughout a logging event and subsequent recovery and found that they constitute a substantial and persistent net carbon source. Consistent with existing literature, our study showed a significantly greater woody biomass gain across moderately and heavily logged forests compared with unlogged forests, but this was counteracted by much larger carbon losses from soil organic matter and deadwood in logged forests. We estimate an average carbon source of 1.75 ± 0.94 Mg C ha-1 yr-1 within moderately logged plots and 5.23 ± 1.23 Mg C ha-1 yr-1 in unsustainably logged and severely degraded plots, with emissions continuing at these rates for at least one-decade post-logging. Our data directly contradict the default assumption that recovering logged and degraded tropical forests are net carbon sinks, implying the amount of carbon being sequestered across the world's tropical forests may be considerably lower than currently estimated.

Major and persistent shifts in below-ground carbon dynamics and soil respiration following logging in tropical forests.

Soil respiration is the largest carbon efflux from the terrestrial ecosystem to the atmosphere, and selective logging influences soil respiration via changes in abiotic (temperature, moisture) and biotic (biomass, productivity, quantity and quality of necromass inputs) drivers. Logged forests are a predominant feature of the tropical forest landscape, their area exceeding that of intact forest. We quantified both total and component (root, mycorrhiza, litter, and soil organic matter, SOM) soil respiration in logged (n = 5) and old-growth (n = 6) forest plots in Malaysian Borneo, a region which is a global hotspot for emission from forest degradation. We constructed a detailed below-ground carbon budget including organic carbon inputs into the system via litterfall and root turnover. Total soil respiration was significantly higher in logged forests than in old-growth forests (14.3 ± 0.23 and 12.7 ± 0.60 Mg C ha-1  year-1 , respectively, p = 0.037). This was mainly due to the higher SOM respiration in logged forests (55 ± 3.1% of the total respiration in logged forests vs. 50 ± 3.0% in old-growth forests). In old-growth forests, annual SOM respiration was equal to the organic carbon inputs into the soil (difference between SOM respiration and inputs 0.18 Mg C ha-1  year-1 , with 90% confidence intervals of -0.41 and 0.74 Mg C ha-1  year-1 ), indicating that the system is in equilibrium, while in logged forests SOM respiration exceeded the inputs by 4.2 Mg C ha-1  year-1 (90% CI of 3.6 and 4.9 Mg C ha-1  year-1 ), indicating that the soil is losing carbon. These results contribute towards understanding the impact of logging on below-ground carbon dynamics, which is one of the key uncertainties in estimating emissions from forest degradation. This study demonstrates how significant perturbation of the below-ground carbon balance, and consequent net soil carbon emissions, can persist for decades after a logging event in tropical forests.

Short- and long-term carbon emissions from oil palm plantations converted from logged tropical peat swamp forest.

Need for regional economic development and global demand for agro-industrial commodities have resulted in large-scale conversion of forested landscapes to industrial agriculture across South East Asia. However, net emissions of CO2 from tropical peatland conversions may be significant and remain poorly quantified, resulting in controversy around the magnitude of carbon release following conversion. Here we present long-term, whole ecosystem monitoring of carbon exchange from two oil palm plantations on converted tropical peat swamp forest. Our sites compare a newly converted oil palm plantation (OPnew) to a mature oil palm plantation (OPmature) and combine them in the context of existing emission factors. Mean annual net emission (NEE) of CO2 measured at OPnew during the conversion period (137.8 Mg CO2  ha-1  year-1 ) was an order of magnitude lower during the measurement period at OPmature (17.5 Mg CO2  ha-1  year-1 ). However, mean water table depth (WTD) was shallower (0.26 m) than a typical drainage target of 0.6 m suggesting our emissions may be a conservative estimate for mature plantations, mean WTD at OPnew was more typical at 0.54 m. Reductions in net emissions were primarily driven by increasing biomass accumulation into highly productive palms. Further analysis suggested annual peat carbon losses of 24.9 Mg CO2 -C ha-1  year-1 over the first 6 years, lower than previous estimates for this early period from subsidence studies, losses reduced to 12.8 Mg CO2 -C ha-1  year-1 in the later, mature phase. Despite reductions in NEE and carbon loss over time, the system remained a large net source of carbon to the atmosphere after 12 years with the remaining 8 years of a typical plantation's rotation unlikely to recoup losses. These results emphasize the need for effective protection of tropical peatlands globally and strengthening of legislative enforcement where moratoria on peatland conversion already exist.

Linking functional traits to multiscale statistics of leaf venation networks.

Leaf venation networks evolved along several functional axes, including resource transport, damage resistance, mechanical strength, and construction cost. Because functions may depend on architectural features at different scales, network architecture may vary across spatial scales to satisfy functional tradeoffs. We develop a framework for quantifying network architecture with multiscale statistics describing elongation ratios, circularity ratios, vein density, and minimum spanning tree ratios. We quantify vein networks for leaves of 260 southeast Asian tree species in samples of up to 2 cm2 , pairing multiscale statistics with traits representing axes of resource transport, damage resistance, mechanical strength, and cost. We show that these multiscale statistics clearly differentiate species' architecture and delineate a phenotype space that shifts at larger scales; functional linkages vary with scale and are weak, with vein density, minimum spanning tree ratio, and circularity ratio linked to mechanical strength (measured by force to punch) and elongation ratio and circularity ratio linked to damage resistance (measured by tannins); and phylogenetic conservatism of network architecture is low but scale-dependent. This work provides tools to quantify the function and evolution of venation networks. Future studies including primary and secondary veins may uncover additional insights.

Imaging spectroscopy reveals the effects of topography and logging on the leaf chemistry of tropical forest canopy trees.

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.

Logging disturbance shifts net primary productivity and its allocation in Bornean tropical forests.

Tropical forests play a major role in the carbon cycle of the terrestrial biosphere. Recent field studies have provided detailed descriptions of the carbon cycle of mature tropical forests, but logged or secondary forests have received much less attention. Here, we report the first measures of total net primary productivity (NPP) and its allocation along a disturbance gradient from old-growth forests to moderately and heavily logged forests in Malaysian Borneo. We measured the main NPP components (woody, fine root and canopy NPP) in old-growth (n = 6) and logged (n = 5) 1 ha forest plots. Overall, the total NPP did not differ between old-growth and logged forest (13.5 ± 0.5 and 15.7 ± 1.5 Mg C ha-1  year-1 respectively). However, logged forests allocated significantly higher fraction into woody NPP at the expense of the canopy NPP (42% and 48% into woody and canopy NPP, respectively, in old-growth forest vs 66% and 23% in logged forest). When controlling for local stand structure, NPP in logged forest stands was 41% higher, and woody NPP was 150% higher than in old-growth stands with similar basal area, but this was offset by structure effects (higher gap frequency and absence of large trees in logged forest). This pattern was not driven by species turnover: the average woody NPP of all species groups within logged forest (pioneers, nonpioneers, species unique to logged plots and species shared with old-growth plots) was similar. Hence, below a threshold of very heavy disturbance, logged forests can exhibit higher NPP and higher allocation to wood; such shifts in carbon cycling persist for decades after the logging event. Given that the majority of tropical forest biome has experienced some degree of logging, our results demonstrate that logging can cause substantial shifts in carbon production and allocation in tropical forests.

Oil palm (Elaeis guineensis) plantation on tropical peatland in South East Asia: Photosynthetic response to soil drainage level for mitigation of soil carbon emissions.

While existing moratoria in Indonesia and Malaysia should preclude continued large-scale expansion of palm oil production into new areas of South-East Asian tropical peatland, existing plantations in the region remain a globally significant source of atmospheric carbon due to drainage driven decomposition of peatland soils. Previous studies have made clear the direct link between drainage depth and peat carbon decomposition and significant reductions in the emission rate of CO2 can be made by raising water tables nearer to the soil surface. However, the impact of such changes on palm fruit yield is not well understood and will be a critical consideration for plantation managers. Here we take advantage of very high frequency, long-term monitoring of canopy-scale carbon exchange at a mature oil palm plantation in Malaysian Borneo to investigate the relationship between drainage level and photosynthetic uptake and consider the confounding effects of light quality and atmospheric vapour pressure deficit. Canopy modelling from our dataset demonstrated that palms were exerting significantly greater stomatal control at deeper water table depths (WTD) and the optimum WTD for photosynthesis was found to be between 0.3 and 0.4 m below the soil surface. Raising WTD to this level, from the industry typical drainage level of 0.6 m, could increase photosynthetic uptake by 3.6 % and reduce soil surface emission of CO2 by 11 %. Our study site further showed that despite being poorly drained compared to other planting blocks at the same plantation, monthly fruit bunch yield was, on average, 14 % greater. While these results are encouraging, and at least suggest that raising WTD closer to the soil surface to reduce emissions is unlikely to produce significant yield penalties, our results are limited to a single study site and more work is urgently needed to confirm these results at other plantations.

Functional susceptibility of tropical forests to climate change.

Tropical forests are some of the most biodiverse ecosystems in the world, yet their functioning is threatened by anthropogenic disturbances and climate change. Global actions to conserve tropical forests could be enhanced by having local knowledge on the forests' functional diversity and functional redundancy as proxies for their capacity to respond to global environmental change. Here we create estimates of plant functional diversity and redundancy across the tropics by combining a dataset of 16 morphological, chemical and photosynthetic plant traits sampled from 2,461 individual trees from 74 sites distributed across four continents together with local climate data for the past half century. Our findings suggest a strong link between climate and functional diversity and redundancy with the three trait groups responding similarly across the tropics and climate gradient. We show that drier tropical forests are overall less functionally diverse than wetter forests and that functional redundancy declines with increasing soil water and vapour pressure deficits. Areas with high functional diversity and high functional redundancy tend to better maintain ecosystem functioning, such as aboveground biomass, after extreme weather events. Our predictions suggest that the lower functional diversity and lower functional redundancy of drier tropical forests, in comparison with wetter forests, may leave them more at risk of shifting towards alternative states in face of further declines in water availability across tropical regions.

An assessment of oil palm plantation aboveground biomass stocks on tropical peat using destructive and non-destructive methods.

The recent expansion of oil palm (OP, Elaeis guineensis) plantations into tropical forest peatlands has resulted in ecosystem carbon emissions. However, estimates of net carbon flux from biomass changes require accurate estimates of the above ground biomass (AGB) accumulation rate of OP on peat. We quantify the AGB stocks of an OP plantation on drained peat in Malaysia from 3 to 12 years after planting using destructive harvests supported by non-destructive surveys of a further 902 palms. Peat specific allometric equations for palm (R2 = 0.92) and frond biomass are developed and contrasted to existing allometries for OP on mineral soils. Allometries are used to upscale AGB estimates to the plantation block-level. Aboveground biomass stocks on peat accumulated at ~6.39 ± 1.12 Mg ha-1 per year in the first 12 years after planting, increasing to ~7.99 ± 0.95 Mg ha-1 yr-1 when a 'perfect' plantation was modelled. High inter-palm and inter-block AGB variability was observed in mature classes as a result of variations in palm leaning and mortality. Validation of the allometries defined and expansion of non-destructive inventories across alternative plantations and age classes on peat would further strengthen our understanding of peat OP AGB accumulation rates.

Leaf venation networks of Bornean trees: images and hand-traced segmentations.

The data set contains images of leaf venation networks obtained from tree species in Malaysian Borneo. The data set contains 726 leaves from 295 species comprising 50 families, sampled from eight forest plots in Sabah. Image extents are approximately 1 × 1 cm, or 50 megapixels. All images contain a region of interest in which all veins have been hand traced. The complete data set includes over 30 billion pixels, of which more than 600 million have been validated by hand tracing. These images are suitable for morphological characterization of these species, as well as for training of machine-learning algorithms that segment biological networks from images. Data are made available under the Open Data Commons Attribution License. You are free to copy, distribute, and use the database; to produce works from the database; and to modify, transform, and build upon the database. You must attribute any public use of the database, or works produced from the database, in the manner specified in the license. For any use or redistribution of the database, or works produced from it, you must make clear to others the license of the database and keep intact any notices on the original database.

Effects of water management and cultivar on carbon dynamics, plant productivity and biomass allocation in European rice systems.

Water saving techniques, such as alternate wetting and drying (AWD), are becoming a necessity in modern rice farming because of climate change mitigation and growing water use scarcity. Reducing water can vastly reduce methane (CH4) emissions; however, this net climate benefit may be offset by enhanced carbon dioxide (CO2) emissions from soil. The main aims of this study were: to determine the effects of AWD on yield and ecosystem C dynamics, and to establish the underlying mechanistic basis for observed trends in net ecosystem C gain or loss in an Italian rice paddy. We investigated the effects of conventional water management (i.e. conventionally flooded paddy; CF) and AWD on biomass accumulation (aboveground, belowground, grain), key ecosystem C fluxes (net ecosystem exchange (NEE), net primary productivity (NPP), gross primary productivity (GPP), ecosystem respiration (ER), autotrophic respiration (RA), heterotrophic respiration (RH)), and soil organic matter (SOM) decay for four common commercial European rice cultivars. The most significant finding was that neither treatment nor cultivar affected NEE, GPP, ER or SOM decomposition. RA was the dominant contributor to ER for both CF and AWD treatments. Cultivar and treatment affected the total biomass of the rice plants; specifically, with greater root production in CF compared to AWD. Importantly, there was no effect of treatment on the overall yield for any cultivar. Possibly, the wetting-drying cycles may have been insufficient to allow substantial soil C metabolism or there was a lack of labile substrate in the soil. These results imply that AWD systems may not be at risk of enhancing soil C loss, making it a viable solution for climate change mitigation and water conservation. Although more studies are needed, the initial outlook for AWD in Europe is positive; with no net loss of soil C from SOM decomposition, whilst also maintaining yield.