Three decades of post-logging tree community recovery in naturally regenerating and actively restored dipterocarp forest in Borneo

Selective logging has affected large areas of tropical forests and there is increasing interest in how to manage selectively logged forests to enhance recovery. However, the impacts of logging and active restoration, by liberation cutting and enrichment planting, on tree community composition are poorly understood compared to trajectories of biomass recovery. Here, we assess the long-term impacts of selective logging and active restoration for biomass recovery on tree species diversity, community composition, and forest structure. We censused all stems ≥2 cm diameter at breast height (DBH) on 46 permanent plots in unlogged, primary forest in the Danum Valley Conservation Area (DVCA; 12 plots, totalling 0.6 ha) and in sites logged 23–35 years prior to the census in the Ulu Segama Forest Reserve adjacent to DVCA (34 plots, totalling 1.7 ha) in Sabah, Malaysian Borneo. Active restoration treatments, including enrichment planting and climber cutting, were implemented on 17 of the logged forest plots 12–24 years prior to the census. Total plot-level basal area and pole (5–10 cm DBH) stem density were lower in logged than unlogged forests, however no difference was found in stem density amongst saplings (2–5 cm DBH) or established trees (≥10 cm DBH). Neither basal area, nor plot-level stem density varied with time since logging at any size class, although sapling and pole stem densities were lower in actively restored than naturally regenerating logged forest. Sapling species diversity was lower in logged than unlogged forest, however there were no other significant effects of logging on tree species richness or diversity indices. Tree species composition, however, differed between logged and unlogged forests across all stem size classes (PERMANOVA), reflected by 23 significant indicator species that were only present in unlogged forest. PERMANOVA tests revealed no evidence that overall species composition changed with time since logging or with active restoration treatments at any size class. However, when naturally regenerating and actively restored communities were compared, two indicator species were identified in naturally regenerating forest and three in actively restored forests. Together our results suggest that selective logging has a lasting effect on tree community composition regardless of active restoration treatments and, even when species richness and diversity are stable, species composition remains distinct from unlogged forest for more than two decades post-harvest. Active restoration efforts should be targeted, monitored, and refined to try to ensure positive outcomes for multiple metrics of forest recovery

Land-use change and propagule pressure promote plant invasions in tropical rainforest remnants

Context: Intact tropical rainforests are considered robust to plant invasions. However, land-use change alters the structure and species composition of native forest, opening up tropical landscapes to invasion. Yet, the relative roles of key drivers on tropical forest invasions remain little investigated. Objectives: We examine factors affecting plant invasion of rainforest remnants in oil-palm dominated landscapes in Sabah, Malaysian Borneo. We hypothesized that invasion is greater in highly fragmented landscapes, and in disturbed forests with lower native plant diversity (cf. old-growth rainforests). Methods: Native and exotic plants were surveyed in 47 plots at 17 forest sites, spanning gradients in landscape-scale fragmentation and local forest disturbance. Using partial least squares path-modelling, we examined correlations between invasion, fragmentation, forest disturbance, propagule pressure, soil characteristics and native plant community. Results: We recorded 6999 individuals from 329 genera in total, including eight exotic species (0–51% of individuals/plot, median = 1.4%) representing shrubs, forbs, graminoids and climbers. The best model (R2 = 0.343) revealed that invasion was correlated with disturbance and propagule pressure (high prevalence of exotic species in plantation matrix), the latter being driven by greater fragmentation of the landscape. Our models revealed a significant negative correlation between invasion and native tree seedlings and sapling community diversity. Conclusions: Increasing landscape fragmentation promotes exotic plant invasion in remnant tropical forests, especially if local disturbance is high. The association between exotic species invasion and young native tree community may have impacts for regeneration given that fragmentation is predicted to increase and so plant invasion may become more prevalent

Trait filtering during exotic plant invasion of tropical rainforest remnants along a disturbance gradient

1. Human‐modified tropical landscapes are often invaded by exotic plant species, but relatively few species are able to colonise remnant areas of rainforest embedded within such landscapes. The functional traits of successful invaders of natural versus anthropogenic habitats are poorly known, especially in tropical regions, and identifying such traits provides insight into the mechanisms that drive invasion. Here, we examine the invasion of tropical rainforest remnants along a disturbance gradient, within a human‐modified agricultural landscape, and determine whether exotic species that invade these forests are selected according to particular traits. 2. We surveyed the occurrence of 18 exotic species along 100‐m transects in four habitats—oil palm road, forest‐oil palm edges and disturbed and intact forest within rainforest remnants—at 21 sites across Sabah, Malaysian Borneo. We collated data on four functional traits relevant to the barriers plants encounter when colonising new environments (e.g. dispersal and persistence) and tested whether trait filtering occurs during invasion of rainforest remnants. 3. Exotic species richness declined significantly from oil palm (mean 9.2 species per transect) to forest edge (7.8 species) to inside rainforest remnants (3.1 species in disturbed forest), and only one species, Clidemia hirta, invaded intact forest. Exotic communities within rainforest remnants had long‐distance (vertebrate) dispersal, were woodier and had taller maximum heights, compared to those found in oil palm. For each trait, the community‐weighted mean for the forest edge community was intermediate between oil palm and disturbed forest, suggesting trait filtering during the invasion of rainforest remnants. 4. Our study provides strong evidence that trait filtering occurs during invasion from human‐modified agricultural habitats into previously disturbed forests via the forest edge. Successful invasion of rainforest remnants requires relatively long‐distance dispersal, in particular by vertebrates, as well as traits that are more similar to those of native forest species (i.e. tall and woody), making these exotic species more able to compete and persist in that environment. Our results show that disturbed tropical rainforests with open canopies are susceptible to invasion and highlight the traits of exotic species which can invade rainforest habitats, and which may pose a threat to regenerating tropical rainforests

Bornean tropical forests recovering from logging at risk of regeneration failure

AUTHOR CONTRIBUTIONS David C. Bartholomew: Conceptualization; formal analysis; investigation; methodology; project administration; visualization; writing – original draft; writing – review and editing. Robin Hayward: Formal analysis; methodology; visualization; writing – original draft; writing – review and editing. David F. R. P. Burslem: Conceptualization; funding acquisition; methodology; supervision; writing – review and editing. Paulo R. L. Bittencourt: Conceptualization; investigation; methodology; writing – review and editing. Daniel Chapman: Formal analysis; writing – review and editing. Mohd. Aminur Faiz Bin Suis: Project administration. Reuben Nilus: Project administration. Michael J. O'Brien: Conceptualization; investigation; methodology; project administration; writing – review and editing. Glen Reynolds: Project administration. Lucy Rowland: Conceptualization; funding acquisition; methodology; supervision; writing – review and editing. Lindsay F. Banin: Conceptualization; funding acquisition; methodology; supervision; writing – review and editing. Daisy Dent: Conceptualization; funding acquisition; methodology; supervision; writing – review and editing.Peer reviewe

Differential nutrient limitation and tree height control leaf physiology, supporting niche partitioning in tropical dipterocarp forests

1. Revealing the mechanisms of environmental niche partitioning within lowland tropical forests is important for understanding the drivers of current species distributions and potential vulnerability to environmental change. Tropical forest structure and species composition change across edaphic gradients in Borneo over short distances. However, our understanding of how edaphic conditions affect tree physiology and whether these relationships drive niche partitioning within Bornean forests remains incomplete. 2. This study evaluated how leaf physiological function changes with nutrient availability across a fine-scale edaphic gradient and whether these relationships vary according to tree height. Furthermore, we tested whether intraspecific leaf trait variation allows generalist species to populate a wider range of environments. 3. We measured leaf traits of 218 trees ranging in height from 4 to 66 m from 13 dipterocarp species within four tropical forest types (alluvial, mudstone, sandstone and kerangas) occurring along an <5 km edaphic gradient in North Borneo. The traits measured included saturating photosynthesis (Asat), maximum photosynthetic capacity (Vcmax), leaf dark respiration (Rleaf), leaf mass per area (LMA), leaf thickness, minimum stomatal conductance (gdark) and leaf nutrient concentrations (N, P, Ca, K and Mg). 4. Across all species, leaf traits varied consistently in response to soil nutrient availability across forest types except Rleaf_mass, [Mg]leaf and [Ca]leaf. Changes in photosynthesis and respiration rates were related to different leaf nutrients across forest types, with greater nutrient-use efficiency in more nutrient-poor environments. Generalist species partially or fully compensated reductions in mass-based photosynthesis through increasing LMA in more nutrient-poor environments. 5. Leaf traits also varied with tree height, except Vcmax_mass, but only in response to height-related modifications of leaf morphology (LMA and leaf thickness). These height–trait relationships did not vary across the edaphic gradient, except for Asat, [N]leaf, [P]leaf and [K]leaf. 6. Our results highlight that modification of leaf physiological function and morphology act as important adaptations for Bornean dipterocarps in response to edaphic and vertical environmental gradients. Meanwhile, multiple nutrients appear to contribute to niche partitioning and could drive species distributions and high biodiversity within Bornean forest landscapes

Making waves. Bridging theory and practice towards multiple stressor management in freshwater ecosystems

Embargo until February 26, 2023Despite advances in conceptual understanding, single-stressor abatement approaches remain common in the management of fresh waters, even though they can produce unexpected ecological responses when multiple stressors interact. Here we identify limitations restricting the development of multiple-stressor management strategies and address these, bridging theory and practice, within a novel empirical framework. Those critical limitations include that (i) monitoring schemes fall short of accounting for theory on relationships between multiple-stressor interactions and ecological responses, (ii) current empirical modelling approaches neglect the prevalence and intensity of multiple-stressor interactions, and (iii) mechanisms of stressor interactions are often poorly understood. We offer practical recommendations for the use of empirical models and experiments to predict the effects of freshwater degradation in response to changes in multiple stressors, demonstrating this approach in a case study. Drawing on our framework, we offer practical recommendations to support the development of effective management strategies in three general multiple-stressor scenarios.acceptedVersio

Fifteen essential science advances needed for effective restoration of the world's forest landscapes

There has never been a more pressing and opportune time for science and practice to collaborate towards restoration of the world's forests. Multiple uncertainties remain for achieving successful, long-term forest landscape restoration (FLR). In this article, we use expert knowledge and literature review to identify knowledge gaps that need closing to advance restoration practice, as an introduction to a landmark theme issue on FLR and the UN Decade on Ecosystem Restoration. Aligned with an Adaptive Management Cycle for FLR, we identify 15 essential science advances required to facilitate FLR success for nature and people. They highlight that the greatest science challenges lie in the conceptualization, planning and assessment stages of restoration, which require an evidence base for why, where and how to restore, at realistic scales. FLR and underlying sciences are complex, requiring spatially explicit approaches across disciplines and sectors, considering multiple objectives, drivers and trade-offs critical for decision-making and financing. The developing tropics are a priority region, where scientists must work with stakeholders across the Adaptive Management Cycle. Clearly communicated scientific evidence for action at the outset of restoration planning will enable donors, decision makers and implementers to develop informed objectives, realistic targets and processes for accountability. This article paves the way for 19 further articles in this theme issue, with author contributions from across the world. This introduction article is part of the theme issue ‘Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration’

Conservation set-asides improve carbon storage and support associated plant diversity in certified sustainable oil palm plantations

Maintaining forest conservation set-asides is a key criterion of sustainability certification of many crops that drive tropical deforestation, but their value for carbon storage and associated biodiversity is unclear. We conducted vegetation measurements to examine the benefits of set-asides for aboveground carbon stocks (AGC) in certified oil palm plantations on Borneo, and whether their AGC is positively associated with plant diversity. The mean estimated AGC of live trees and palms ≥10 cm diameter in set-asides in certified oil palm plantations (52.8 Mg ha−1) was >1.5-times that of oil palm (30.3 Mg ha−1), with some plots supporting similar AGC to primary forest. For lowland Borneo, we estimate that the average AGC of oil palm plantations with 10% coverage of set-asides is up to 20% greater than the average AGC of oil palm plantations without set-asides, newly demonstrating carbon storage as a benefit of conservation set-asides. We found positive relationships between AGC and plant diversity, highlighting the carbon–biodiversity co-benefits of set-asides. However, set-asides had a lower density of tree seedlings than continuous primary forest, indicating potential suppression of future tree regeneration and AGC. Our findings support the application of zero-deforestation during agricultural development, to conserve areas of remaining forest with high AGC and high biodiversity. We recommend management practices that boost regeneration in existing set-asides (e.g. enrichment planting), which would be most effective in larger set-asides, and could substantially increase the AGC of agricultural landscapes without removing land from production, and help conserve forest-dependent biodiversity

Restoring understory and riparian areas in oil palm plantations does not increase greenhouse gas fluxes

Oil palm (OP) plantations have replaced large areas of forest in the tropical landscape of Southeast Asia and are major emitters of greenhouse gases (GHGs). To move towards more environmentally friendly plantation management, a hopeful approach is to implement strategies to increase vegetation complexity. These options include relaxed management of understory vegetation to increase complexity in productive plantations, passive restoration of forest areas around rivers by leaving mature oil palm during replanting, and active forest restoration along river margins with planting of forest trees. These practices have the potential to deliver a range of benefits such as soil protection, reduced erosion and sedimentation in rivers, pest control and support for biodiversity, but little is known about their impact on greenhouse gas fluxes. The aim of this study was to assess the impact of improved understory growth management and the use of riparian forestry on GHG fluxes in OP plantations, making use of two long-term experiments (the Biodiversity and Ecosystem Function in Tropical Agriculture Understory Vegetation (BEFTA UV) Project; the Riparian Ecosystem Restoration in Tropical Agriculture (RERTA) Project) in Riau Province, Sumatra, Indonesia. We measured nitrous oxide (N2O), methane (CH4) and ecosystem respiration (CO2) from mature OP sites with different levels of understory vegetation and different riparian buffer restoration treatments using the static chamber method. We used linear mixed effects models to test for treatment effects, whilst accounting for soil moisture and experimental design factors (time and space). The understory vegetation treatments (normal, reduced and enhanced complexity of understory) had no effect on N2O and CH4 flux. Regarding differences in ecosystem respiration, effects attributable to the understory vegetation treatments were not strong. For the riparian restoration treatments, the fixed effects variables in the models explained little variation in the fluxes of all GHGs. Therefore, given the proven benefits of more complex understory vegetation for supporting biodiversity and healthy ecosystem functioning, plus the potential for restored riparian buffers to support biodiversity and services and to reduce GHG emissions over time, our findings reinforce the concept that these features bring environmental benefits in OP landscapes, with no measurable effects on GHG emissions