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Active restoration through silvicultural treatments (enrichment planting, cutting climbers and liberation thinning) is considered an important intervention in logged forests. However, its ability to enhance regeneration is key for long-term recovery of logged forests, which remains poorly understood, particularly for the production and survival of seedlings in subsequent generations. To understand the long-term impacts of logging and restoration we tracked the diversity, survival and traits of seedlings that germinated immediately after a mast fruiting in North Borneo in unlogged and logged forests 30–35 years after logging. We monitored 5119 seedlings from germination for ~1.5 years across a mixed landscape of unlogged forests (ULs), naturally regenerating logged forests (NR) and actively restored logged forests via rehabilitative silvicultural treatments (AR), 15–27 years after restoration. We measured 14 leaf, root and biomass allocation traits on 399 seedlings from 15 species. Soon after fruiting, UL and AR forests had higher seedling densities than NR forest, but survival was the lowest in AR forests in the first 6 months. Community composition differed among forest types; AR and NR forests had lower species richness and lower evenness than UL forests by 5–6 months post-mast but did not differ between them. Differences in community composition altered community-weighted mean trait values across forest types, with higher root biomass allocation in NR relative to UL forest. Traits influenced mortality ~3 months post-mast, with more acquisitive traits and relative aboveground investment favoured in AR forests relative to UL forests. Our findings of reduced seedling survival and diversity suggest long time lags in post-logging recruitment, particularly for some taxa. Active restoration of logged forests recovers initial seedling production, but elevated mortality in AR forests lowers the efficacy of active restoration to enhance recruitment or diversity of seedling

Active restoration through silvicultural treatments (enrichment planting, cutting climbers and liberation thinning) is considered an important intervention in logged forests. However, its ability to enhance regeneration is key for long-term recovery of logged forests, which remains poorly understood, particularly for the production and survival of seedlings in subsequent generations. To understand the long-term impacts of logging and restoration we tracked the diversity, survival and traits of seedlings that germinated immediately after a mast fruiting in North Borneo in unlogged and logged forests 30–35 years after logging. We monitored 5119 seedlings from germination for ~1.5 years across a mixed landscape of unlogged forests (ULs), naturally regenerating logged forests (NR) and actively restored logged forests via rehabilitative silvicultural treatments (AR), 15–27 years after restoration. We measured 14 leaf, root and biomass allocation traits on 399 seedlings from 15 species. Soon after fruiting, UL and AR forests had higher seedling densities than NR forest, but survival was the lowest in AR forests in the first 6 months. Community composition differed among forest types; AR and NR forests had lower species richness and lower evenness than UL forests by 5–6 months post-mast but did not differ between them. Differences in community composition altered community-weighted mean trait values across forest types, with higher root biomass allocation in NR relative to UL forest. Traits influenced mortality ~3 months post-mast, with more acquisitive traits and relative aboveground investment favoured in AR forests relative to UL forests. Our findings of reduced seedling survival and diversity suggest long time lags in post-logging recruitment, particularly for some taxa. Active restoration of logged forests recovers initial seedling production, but elevated mortality in AR forests lowers the efficacy of active restoration to enhance recruitment or diversity of seedling

Active restoration through silvicultural treatments (enrichment planting, cutting climbers and liberation thinning) is considered an important intervention in logged forests. However, its ability to enhance regeneration is key for long-term recovery of logged forests, which remains poorly understood, particularly for the production and survival of seedlings in subsequent generations. To understand the long-term impacts of logging and restoration we tracked the diversity, survival and traits of seedlings that germinated immediately after a mast fruiting in North Borneo in unlogged and logged forests 30–35 years after logging. We monitored 5119 seedlings from germination for ~1.5 years across a mixed landscape of unlogged forests (ULs), naturally regenerating logged forests (NR) and actively restored logged forests via rehabilitative silvicultural treatments (AR), 15–27 years after restoration. We measured 14 leaf, root and biomass allocation traits on 399 seedlings from 15 species. Soon after fruiting, UL and AR forests had higher seedling densities than NR forest, but survival was the lowest in AR forests in the first 6 months. Community composition differed among forest types; AR and NR forests had lower species richness and lower evenness than UL forests by 5–6 months post-mast but did not differ between them. Differences in community composition altered community-weighted mean trait values across forest types, with higher root biomass allocation in NR relative to UL forest. Traits influenced mortality ~3 months post-mast, with more acquisitive traits and relative aboveground investment favoured in AR forests relative to UL forests. Our findings of reduced seedling survival and diversity suggest long time lags in post-logging recruitment, particularly for some taxa. Active restoration of logged forests recovers initial seedling production, but elevated mortality in AR forests lowers the efficacy of active restoration to enhance recruitment or diversity of seedling

Active restoration through silvicultural treatments (enrichment planting, cutting climbers and liberation thinning) is considered an important intervention in logged forests. However, its ability to enhance regeneration is key for long-term recovery of logged forests, which remains poorly understood, particularly for the production and survival of seedlings in subsequent generations. To understand the long-term impacts of logging and restoration we tracked the diversity, survival and traits of seedlings that germinated immediately after a mast fruiting in North Borneo in unlogged and logged forests 30–35 years after logging. We monitored 5119 seedlings from germination for ~1.5 years across a mixed landscape of unlogged forests (ULs), naturally regenerating logged forests (NR) and actively restored logged forests via rehabilitative silvicultural treatments (AR), 15–27 years after restoration. We measured 14 leaf, root and biomass allocation traits on 399 seedlings from 15 species. Soon after fruiting, UL and AR forests had higher seedling densities than NR forest, but survival was the lowest in AR forests in the first 6 months. Community composition differed among forest types; AR and NR forests had lower species richness and lower evenness than UL forests by 5–6 months post-mast but did not differ between them. Differences in community composition altered community-weighted mean trait values across forest types, with higher root biomass allocation in NR relative to UL forest. Traits influenced mortality ~3 months post-mast, with more acquisitive traits and relative aboveground investment favoured in AR forests relative to UL forests. Our findings of reduced seedling survival and diversity suggest long time lags in post-logging recruitment, particularly for some taxa. Active restoration of logged forests recovers initial seedling production, but elevated mortality in AR forests lowers the efficacy of active restoration to enhance recruitment or diversity of seedling

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 thickne

Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

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 thickne

Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

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 thickne

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 thickne

Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

Fine-scale topographic–edaphic gradients are common in tropical forests and drive species spatial turnover and marked changes in forest structure and function. We evaluate how hydraulic traits of tropical tree species relate to vertical and horizontal spatial niche specialization along such a gradient. Along a topographic–edaphic gradient with uniform climate in Borneo, we measured six key hydraulic traits in 156 individuals of differing heights in 13 species of Dipterocarpaceae. We investigated how hydraulic traits relate to habitat, tree height and their interaction on this gradient. Embolism resistance increased in trees on sandy soils but did not vary with tree height. By contrast, water transport capacity increased on sandier soils and with increasing tree height. Habitat and height only interact for hydraulic efficiency, with slope for height changing from positive to negative from the clay-rich to the sandier soil. Habitat type influenced trait–trait relationships for all traits except wood density. Our data reveal that variation in the hydraulic traits of dipterocarps is driven by a combination of topographic–edaphic conditions, tree height and taxonomic identity. Our work indicates that hydraulic traits play a significant role in shaping forest structure across topographic–edaphic and vertical gradients and may contribute to niche specialization among dipterocarp species.

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 thickne