Balancing economic and ecological functions in smallholder and industrial oil palm plantations.

The expansion of the oil palm industry in Indonesia has improved livelihoods in rural communities, but comes at the cost of biodiversity and ecosystem degradation. Here, we investigated ways to balance ecological and economic outcomes of oil palm cultivation. We compared a wide range of production systems, including smallholder plantations, industrialized company estates, estates with improved agronomic management, and estates with native tree enrichment. Across all management types, we assessed multiple indicators of biodiversity, ecosystem functions, management, and landscape structure to identify factors that facilitate economic-ecological win-wins, using palm yields as measure of economic performance. Although, we found that yields in industrialized estates were, on average, twice as high as those in smallholder plantations, ecological indicators displayed substantial variability across systems, regardless of yield variations, highlighting potential for economic-ecological win-wins. Reducing management intensity (e.g., mechanical weeding instead of herbicide application) did not lower yields but improved ecological outcomes at moderate costs, making it a potential measure for balancing economic and ecological demands. Additionally, maintaining forest cover in the landscape generally enhanced local biodiversity and ecosystem functioning within plantations. Enriching plantations with native trees is also a promising strategy to increase ecological value without reducing productivity. Overall, we recommend closing yield gaps in smallholder cultivation through careful intensification, whereas conventional plantations could reduce management intensity without sacrificing yield. Our study highlights various pathways to reconcile the economics and ecology of palm oil production and identifies management practices for a more sustainable future of oil palm cultivation.

Effects of plant diversity on productivity strengthen over time due to trait-dependent shifts in species overyielding.

Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

Transformation scenarios towards multifunctional landscapes: A multi-criteria land-use allocation model applied to Jambi Province, Indonesia.

In tropical regions, shifting from forests and traditional agroforestry to intensive plantations generates conflicts between human welfare (farmers' demands and societal needs) and environmental protection. Achieving sustainability in this transformation will inevitably involve trade-offs between multiple ecological and socioeconomic functions. To address these trade-offs, our study used a new methodological approach allowing the identification of transformation scenarios, including theoretical landscape compositions that satisfy multiple ecological functions (i.e., structural complexity, microclimatic conditions, organic carbon in plant biomass, soil organic carbon and nutrient leaching losses), and farmers needs (i.e., labor and input requirements, total income to land, and return to land and labor) while accounting for the uncertain provision of these functions and having an actual potential for adoption by farmers. We combined a robust, multi-objective optimization approach with an iterative search algorithm allowing the identification of ecological and socioeconomic functions that best explain current land-use decisions. The model then optimized the theoretical land-use composition that satisfied multiple ecological and socioeconomic functions. Between these ends, we simulated transformation scenarios reflecting the transition from current land-use composition towards a normative multifunctional optimum. These transformation scenarios involve increasing the number of optimized socioeconomic or ecological functions, leading to higher functional richness (i.e., number of functions). We applied this method to smallholder farms in the Jambi Province, Indonesia, where traditional rubber agroforestry, rubber plantations, and oil palm plantations are the main land-use systems. Given the currently practiced land-use systems, our study revealed short-term returns to land as the principal factor in explaining current land-use decisions. Fostering an alternative composition that satisfies additional socioeconomic functions would require minor changes ("low-hanging fruits"). However, satisfying even a single ecological indicator (e.g., reduction of nutrient leaching losses) would demand substantial changes in the current land-use composition ("moonshot"). This would inevitably lead to a profit decline, underscoring the need for incentives if the societal goal is to establish multifunctional agricultural landscapes. With many oil palm plantations nearing the end of their production cycles in the Jambi province, there is a unique window of opportunity to transform agricultural landscapes.

Complementary ecosystem services from multiple land uses highlight the importance of tropical mosaic landscapes.

Tropical agricultural landscapes often consist of a mosaic of different land uses, yet little is known about the spectrum of ecosystem service bundles and materials they provide to rural households. We interviewed 320 households on the different benefits received from prevalent land-use types in north-eastern Madagascar (old-growth forests, forest fragments, vanilla agroforests, woody fallows, herbaceous fallows, and rice paddies) in terms of ecosystem services and plant uses. Old-growth forests and forest fragments were reported as important for regulating services (e.g. water regulation), whilst fallow lands and vanilla agroforests as important for provisioning services (food, medicine, fodder). Households reported the usage of 285 plant species (56% non-endemics) and collected plants from woody fallows for varying purposes, whilst plants from forest fragments, predominantly endemics, were used for construction and weaving. Multiple land-use types are thus complementary for providing ecosystem services, with fallow lands being particularly important. Hence, balancing societal needs and conservation goals should be based on diversified and comprehensive land management.

Partitioning of respired CO2 in newly sprouted Moso bamboo culms.

Stem respiration (R s) plays a vital role in ecosystem carbon cycling. However, the measured efflux on the stem surface (E s) is not always in situ R s but only part of it. A previously proposed mass balance framework (MBF) attempted to explore the multiple partitioning pathways of R s, including sap-flow-transported and internal storage of R s, in addition to E s. This study proposed stem photosynthesis as an additional partitioning pathway to the MBF. Correspondingly, a double-chamber apparatus was designed and applied on newly sprouted Moso bamboo (Phyllostachys edulis) in leafless and leaved stages. R s of newly sprouted bamboo were twice as high in the leafless stage (7.41 ± 2.66 µmol m-2 s-1) than in the leaved stage (3.47 ± 2.43 µmol m-2 s-1). E s accounted for ~80% of R s, while sap flow may take away ~2% of R s in both leafless and leaved stages. Culm photosynthesis accounted for ~9% and 13% of R s, respectively. Carbon sequestration from culm photosynthesis accounted for approximately 2% of the aboveground bamboo biomass in the leafless stage. High culm photosynthesis but low sap flow during the leafless stage and vice versa during the leaved stage make bamboo an outstanding choice for exploring the MBF.

Tree islands enhance biodiversity and functioning in oil palm landscapes.

In the United Nations Decade on Ecosystem Restoration1, large knowledge gaps persist on how to increase biodiversity and ecosystem functioning in cash crop-dominated tropical landscapes2. Here, we present findings from a large-scale, 5-year ecosystem restoration experiment in an oil palm landscape enriched with 52 tree islands, encompassing assessments of ten indicators of biodiversity and 19 indicators of ecosystem functioning. Overall, indicators of biodiversity and ecosystem functioning, as well as multidiversity and ecosystem multifunctionality, were higher in tree islands compared to conventionally managed oil palm. Larger tree islands led to larger gains in multidiversity through changes in vegetation structure. Furthermore, tree enrichment did not decrease landscape-scale oil palm yield. Our results demonstrate that enriching oil palm-dominated landscapes with tree islands is a promising ecological restoration strategy, yet should not replace the protection of remaining forests.

Tree identity and canopy openness mediate oil palm biodiversity enrichment effects on insect herbivory and pollination.

As the extent of oil palm (Elaeis guineensis) cultivation has expanded at the expense of tropical rainforests, enriching conventional large-scale oil palm plantations with native trees has been proposed as a strategy for restoring biodiversity and ecosystem function. However, how tree enrichment affects insect-mediated ecosystem functions is unknown. We investigated impacts on insect herbivory and pollination in the fourth year of a plantation-scale, long-term oil palm biodiversity enrichment experiment in Jambi, Sumatra, Indonesia. Within 48 plots systematically varying in size (25-1600 m2 ) and planted tree species richness (one to six species), we collected response data on vegetation structure, understory insect abundances, and pollinator and herbivore activity on chili plants (Capsicum annuum), which served as indicators of insect-mediated ecosystem functions. We examined the independent effects of plot size, tree species richness, and tree identity on these response variables, using the linear model for random partitions design. The experimental treatments were most associated with vegetation structure: tree identity mattered, as the species Peronema canescens strongly decreased (by approximately one standard deviation) both canopy openness and understory vegetation cover; whereas tree richness only decreased understory flower density. Further, the smallest plots had the lowest understory flower density and richness, presumably because of lower light availability and colonization rates, respectively. Enrichment influenced herbivorous insects and natural enemies in the understory to a lesser extent: both groups had higher abundances in plots with two enrichment species planted, possibly because higher associated tree mortality created more habitat, while herbivores decreased with increasing tree species richness, in line with the resource concentration hypothesis. Linking relationships in structural equation models showed that the negative association between P. canescens and understory vegetation cover was mediated through canopy openness. Likewise, canopy openness mediated increases in herbivore and pollinator insect abundances. Higher pollinator visitation increased phytometer yield, while impacts of insect herbivores on yield were not apparent. Our results demonstrate that even at an early stage, different levels of ecological restoration influence insect-mediated ecosystem functions, mainly through canopy openness. These findings suggest that maintaining some canopy gaps while enrichment plots develop may be beneficial for increasing habitat heterogeneity and insect-mediated ecosystem functions.

Win-win opportunities combining high yields with high multi-taxa biodiversity in tropical agroforestry.

Resolving ecological-economic trade-offs between biodiversity and yields is a key challenge when addressing the biodiversity crisis in tropical agricultural landscapes. Here, we focused on the relation between seven different taxa (trees, herbaceous plants, birds, amphibians, reptiles, butterflies, and ants) and yields in vanilla agroforests in Madagascar. Agroforests established in forests supported overall 23% fewer species and 47% fewer endemic species than old-growth forests, and 14% fewer endemic species than forest fragments. In contrast, agroforests established on fallows had overall 12% more species and 38% more endemic species than fallows. While yields increased with vanilla vine density and length, non-yield related variables largely determined biodiversity. Nonetheless, trade-offs existed between yields and butterflies as well as  reptiles. Vanilla yields were generally unrelated to richness of trees, herbaceous plants, birds, amphibians, reptiles, and ants, opening up possibilities for conservation outside of protected areas and restoring degraded land to benefit farmers and biodiversity alike.

Land-use trajectories for sustainable land system transformations: Identifying leverage points in a global biodiversity hotspot.

Sustainable land-system transformations are necessary to avert biodiversity and climate collapse. However, it remains unclear where entry points for transformations exist in complex land systems. Here, we conceptualize land systems along land-use trajectories, which allows us to identify and evaluate leverage points, i.e., entry points on the trajectory where targeted interventions have particular leverage to influence land-use decisions. We apply this framework in the biodiversity hotspot Madagascar. In the northeast, smallholder agriculture results in a land-use trajectory originating in old-growth forests and spanning from forest fragments to shifting hill rice cultivation and vanilla agroforests. Integrating interdisciplinary empirical data on seven taxa, five ecosystem services, and three measures of agricultural productivity, we assess trade-offs and cobenefits of land-use decisions at three leverage points along the trajectory. These trade-offs and cobenefits differ between leverage points: Two leverage points are situated at the conversion of old-growth forests and forest fragments to shifting cultivation and agroforestry, resulting in considerable trade-offs, especially between endemic biodiversity and agricultural productivity. Here, interventions enabling smallholders to conserve forests are necessary. This is urgent since ongoing forest loss threatens to eliminate these leverage points due to path dependency. The third leverage point allows for the restoration of land under shifting cultivation through vanilla agroforests and offers cobenefits between restoration goals and agricultural productivity. The co-occurring leverage points highlight that conservation and restoration are simultaneously necessary to avert collapse of multifunctional mosaic landscapes. Methodologically, the framework highlights the importance of considering path dependency along trajectories to achieve sustainable land-system transformations.

Dataset on microclimate and drone-based thermal patterns within an oil palm agroforestry system.

Microclimate and Land Surface Temperature (LST) are important analytical variables used to understand complex oil palm agroforestry systems and their effects on biodiversity and ecosystem functions. In order to examine experimental effects of tree species richness (0, 1, 2, 3 or 6), plot size (25 m2, 100 m2, 400 m2, 1600 m2) and stand structural complexity on microclimate and Land Surface Temperature, related data were collected following a strict design. The experiment was carried out in the Jambi province, in Sumatra (Indonesia), as part of the collaborative project EFForTS [Ecological and Socioeconomic Functions of Tropical Lowland Rainforest Transformation Systems]. Microclimate data collected using miniaturized data loggers combined with drone-based thermal data were considered within an oil palm plantation enriched with six target tree species. The timeframe considered for data analysis was 20th September 2017 to 26th September 2017. The experiment data can be used for comparison with data from conventional oil palm agroforestry systems in the tropics. They can more specifically be used as reference to assess microclimate and Land Surface Temperature patterns within similar agroforestry systems.

"Diminishing returns" for leaves of five age-groups of Phyllostachys edulis culms.

PREMISE: Leaf mass (M) and lamina surface area (A) are important functional traits reported to obey a scaling relationship called "diminishing returns" (i.e., M ∝ Aα>1 ). Previous studies have focused primarily on eudicots and ignored whether the age of leaves affects the numerical value of the scaling exponent (i.e., α). METHODS: The effect of age was examined using 1623 Phyllostachys edulis leaves from culms differing in age collected in Nanjing, China. The scaling relationships among leaf A, fresh mass (FM), and dry mass (DM) were evaluated using reduced major axis protocols. The bootstrap percentile method was used to test the significance of differences in α-values. RESULTS: Overall, the numerical values of α exceeded 1.0. The scaling relationship between FM and A was statistically more robust than that between DM and A. The scaling exponents of FM vs. A exhibited a "high-low-high-low-high" numerical trend from the oldest to the youngest age-group. FM increased linearly as culm age decreased; the leaf DM per unit area (LMA) exhibited a parabolic trend across the age-groups. CONCLUSIONS: "Diminishing returns" is confirmed for all but one age-group of an important monocot species. The relationship between FM and A was statistically more robust than that between DM and A for each age-group. The FM per unit A decreased with increasing age-groups, whereas the middle age-groups had a greater LMA than the oldest and youngest age-groups. These data are the first to show that the age of shoots affects the scaling relationship between leaf mass and area.

Spring Leafing Phenology Favors Younger Culms of Moso Bamboo: Aspects From Water Use Relations.

As the most widely distributed giant running bamboo species in China, Moso bamboo (Phyllostachys edulis) can accomplish both development of newly sprouted culms and leaf renewal of odd-year-old culms within a few months in spring. The two phenological events in spring may together change water distribution among culms in different age categories within a stand, which may differ from our conventional understanding of the negative age effect on bamboo water use. Therefore, to explore the effect of spring shooting and leaf phenology on age-specific water use of Moso bamboo and potential water redistribution, we monitored water use of four culm age categories (newly sprouted, 1-, 2-, and 3-year-old; namely A0, A1, A2, A3) in spring from March to June 2018. For newly sprouting culms, the spring phenological period was classified into five stages (incubation, culm-elongation, branch-development, leafing, established). Over these phenological stages, age-specific accumulated sap flux density showed different patterns. The oldest culms, A3, were not influenced by leaf renewal and kept nearly constant and less water use than the other aged culms. However, A2, which did not renew their leaves, had the most water use at the two initial stages (incubation, culm-elongation) but consumed less water than A0 and A1 after the fourth stage (leafing). At the end of June, water use of the four age categories sorted in order of A0 > A1 > A2 > A3, which confirms the conventional thought and observations, i.e., a negative age effect. The results indicate that new leaf flushing may benefit younger culms (A1 and A0) more than older culms (A2 and A3), i.e., increasing their transpiration response to radiation and share of the stand transpiration. With the underground connected rhizome system, the bamboo stand as an integration seems to balance its water use among culms of different ages to support the water use of freshly sprouted culms during their developing period.

Trade-offs between multifunctionality and profit in tropical smallholder landscapes.

Land-use transitions can enhance the livelihoods of smallholder farmers but potential economic-ecological trade-offs remain poorly understood. Here, we present an interdisciplinary study of the environmental, social and economic consequences of land-use transitions in a tropical smallholder landscape on Sumatra, Indonesia. We find widespread biodiversity-profit trade-offs resulting from land-use transitions from forest and agroforestry systems to rubber and oil palm monocultures, for 26,894 aboveground and belowground species and whole-ecosystem multidiversity. Despite variation between ecosystem functions, profit gains come at the expense of ecosystem multifunctionality, indicating far-reaching ecosystem deterioration. We identify landscape compositions that can mitigate trade-offs under optimal land-use allocation but also show that intensive monocultures always lead to higher profits. These findings suggest that, to reduce losses in biodiversity and ecosystem functioning, changes in economic incentive structures through well-designed policies are urgently needed.

How a measure of tree structural complexity relates to architectural benefit-to-cost ratio, light availability, and growth of trees.

Aboveground tree architecture is neither fully deterministic nor random. It is likely the result of mechanisms that balance static requirements and light-capturing efficiency. Here, we used terrestrial laser scanning data to investigate the relationship between tree architecture, here addressed using the box-dimension (D b), and the architectural benefit-to-cost ratio, the light availability, and the growth of trees. We detected a clear relationship between D b and the benefit-to-cost ratio for the tested three temperate forest tree species (Fagus sylvatica L., Fraxinus excelsior L., and Acer pseudoplatanus L.). In addition, we could also show that D b is positively related to the growth performance of several tropical tree species. Finally, we observed a negative relationship between the strength of competition enforced on red oak (Quercus rubra L.) trees and their D b. We therefore argue that D b is a meaningful and integrative measure that describes the structural complexity of the aboveground compartments of a plant as well as its relation to structural efficiency (benefit-to-cost ratio), productivity, and growing conditions (competition or availability of light).

Water Transfer Between Bamboo Culms in the Period of Sprouting.

Bamboo culms are connected to neighboring culms via rhizomes, which enable resource exchange between culms. We assessed water transfer between established and neighboring, freshly sprouted culms by thermal dissipation probes (TDP) inserted into culms and the connecting rhizome. During the early phase of sprouting, highest sap flux densities in freshly sprouted culms were observed at night, whereas neighboring established culms had high sap flux densities during daytime. After leaf flushing on freshly sprouted culms, the nighttime peaks disappeared and culms switched to the diurnal sap flux patterns with daytime maxima as observed in established culms. TDP in rhizomes indicated water flowing from the established to the freshly sprouted culms. When the established culms of a clump were cut, freshly sprouted culms without leaves reduced sap flux densities rates by 79%. Our findings thus suggest that bamboos exchange water via rhizomes and that nighttime fluxes are highly important for the support of freshly sprouted culms. The (water) resource support may facilitate the very fast growth of the bamboo shoots, and enable the colonizing of new places.

Tree Water Use Patterns as Influenced by Phenology in a Dry Forest of Southern Ecuador.

Tropical dry forests are composed of tree species with different drought coping strategies and encompass heterogeneous site conditions. Actual water use will be controlled by soil moisture availability. In a premontane dry forest of southern Ecuador, tree water use patterns of four tree species of different phenologies were studied along an elevational gradient, in which soil moisture availability increases with altitude. Main interest was the influence of variation in soil moisture, vapor pressure deficit, species (representing phenology), elevation, and tree diameter on water use. Special emphasis was put on the stem succulent, deciduous Ceiba trichistandra, as high water use rates and drought coping involving stem succulence was to be expected. Tree water use rates increased linearly with diameter across species at high soil water content. However, when soil moisture declined, sap flux densities of the species responded differently. The stem succulent, deciduous Ceiba and other deciduous tree species reduced sap flux sensitively, whereas sap flux densities of the evergreen (broad leaved) Capparis scabrida were increasing. This was also reflected in diurnal hysteresis loops of sap flux vs. vapor pressure deficit (VPD) of the air. Under dry soil conditions, Ceiba and other deciduous tree species had much smaller areas in the hysteresis loop, whereas the area of Capparis was largely enhanced compared to wet conditions. The evergreen Capparis potentially had access to deeper soil water resources as water use patterns suggest that top soil drought was tolerated. The deciduous species followed a drought avoidance strategy by being leafless in the dry season. The stem succulent deciduous Ceiba flushed leaves at the end of the dry season before the rainy season began and also re-flushed early in the dry season after a rain event; however, water use rates at this occasion remained low. Ceiba was also ready for fast and strong response in water use when conditions were most favorable during the wet season. The study thus indicates a strong influence of species' drought coping strategy on water use patterns in tropical dry forests.

Oil Palm and Rubber Tree Water Use Patterns: Effects of Topography and Flooding.

Oil palm and rubber plantations extend over large areas and encompass heterogeneous site conditions. In periods of high rainfall, plants in valleys and at riparian sites are more prone to flooding than plants at elevated topographic positions. We asked to what extent topographic position and flooding affect oil palm and rubber tree water use patterns and thereby influence spatial and temporal heterogeneity of transpiration. In an undulating terrain in the lowlands of Jambi, Indonesia, plantations of the two species were studied in plot pairs consisting of upland and adjacent valley plots. All upland plots were non-flooded, whereas the corresponding valley plots included non-flooded, long-term flooded, and short-term flooded conditions. Within each plot pair, sap flux densities in palms or trees were monitored simultaneously with thermal dissipation probes. In plot pairs with non-flooded valleys, sap flux densities of oil palms were only slightly different between the topographic positions, whereas sap flux densities of rubber trees were higher in the valley than at the according upland site. In pairs with long-term flooded valleys, sap flux densities in valleys were lower than at upland plots for both species, but the reduction was far less pronounced in oil palms than in rubber trees (-22 and -45% in maximum sap flux density, respectively). At these long-term flooded valley plots palm and tree water use also responded less sensitively to fluctuations in micrometeorological variables than at upland plots. In short-term flooded valley plots, sap flux densities of oil palm were hardly affected by flooding, but sap flux densities of rubber trees were reduced considerably. Topographic position and flooding thus affected water use patterns in both oil palms and rubber trees, but the changes in rubber trees were much more pronounced: compared to non-flooded upland sites, the different flooding conditions at valley sites amplified the observed heterogeneity of plot mean water use by a factor of 2.4 in oil palm and by a factor of 4.2 in rubber plantations. Such strong differences between species as well as the pronounced heterogeneity of water use across space and time may be of relevance for eco-hydrological assessments of tropical plantation landscapes.

Spatial and Temporal Localization of Flavonoid Metabolites in Strawberry Fruit (Fragaria × ananassa).

Flavonoids are important metabolites in strawberries (Fragaria × ananassa) because they accomplish an extensive collection of physiological functions and are valuable for human health. However, their localization within the fruit tissue has not been extensively explored. Matrix-assisted laser desorption/ionization mass spectrometric imaging (MALDI-MSI) was employed to shed light on the spatial distribution of flavonoids during fruit development. One wild-type (WT) and two transgenic lines were compared, wherein the transgenic enzymes anthocyanidin reductase (ANRi) and flavonol synthase (FLSi), respectively, were down-regulated using an RNAi-based silencing approach. In most cases, fruit development led to a reduction of the investigated flavonoids in the fruit tissue; as a consequence, they were exclusively present in the skin of mature red fruits. In the case of (epi)catechin dimer, both the ANRi and the WT phenotypes revealed low levels in mature red fruits, whereas the ANRi line bore the lowest relative concentration, as analyzed by liquid chromatography-electrospray ionization multiple-step mass spectrometry (LC-ESI-MSn).

Bioactive secondary metabolites with multiple activities from a fungal endophyte.

In order to replace particularly biohazardous nematocides, there is a strong drive to finding natural product-based alternatives with the aim of containing nematode pests in agriculture. The metabolites produced by the fungal endophyte Fusarium oxysporum 162 when cultivated on rice media were isolated and their structures elucidated. Eleven compounds were obtained, of which six were isolated from a Fusarium spp. for the first time. The three most potent nematode-antagonistic compounds, 4-hydroxybenzoic acid, indole-3-acetic acid (IAA) and gibepyrone D had LC50 values of 104, 117 and 134 µg ml-1 , respectively, after 72 h. IAA is a well-known phytohormone that plays a role in triggering plant resistance, thus suggesting a dual activity, either directly, by killing or compromising nematodes, or indirectly, by inducing defence mechanisms against pathogens (nematodes) in plants. Such compounds may serve as important leads in the development of novel, environmental friendly, nematocides.