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.

Logged tropical forests have amplified and diverse ecosystem energetics.

Old-growth tropical forests are widely recognized as being immensely important for their biodiversity and high biomass1. Conversely, logged tropical forests are usually characterized as degraded ecosystems2. However, whether logging results in a degradation in ecosystem functions is less clear: shifts in the strength and resilience of key ecosystem processes in large suites of species have rarely been assessed in an ecologically integrated and quantitative framework. Here we adopt an ecosystem energetics lens to gain new insight into the impacts of tropical forest disturbance on a key integrative aspect of ecological function: food pathways and community structure of birds and mammals. We focus on a gradient spanning old-growth and logged forests and oil palm plantations in Borneo. In logged forest there is a 2.5-fold increase in total resource consumption by both birds and mammals compared to that in old-growth forests, probably driven by greater resource accessibility and vegetation palatability. Most principal energetic pathways maintain high species diversity and redundancy, implying maintained resilience. Conversion of logged forest into oil palm plantation results in the collapse of most energetic pathways. Far from being degraded ecosystems, even heavily logged forests can be vibrant and diverse ecosystems with enhanced levels of ecological function.

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.

Bornean tropical forests recovering from logging at risk of regeneration failure.

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 communities. This suggests current active restoration practices may fail to overcome barriers to regeneration in logged forests, which may drive long-term changes in future forest plant communities.

A restauração ativa por meio de tratamentos silviculturais (plantio de enriquecimento, corte de trepadeiras e desbaste) é considerada uma intervenção importante em florestas com exploração de madeira. No entanto, sua capacidade de melhorar a regeneração, essencial para a recuperação de longo prazo das florestas exploradas, permanece pouco compreendida, especialmente no que diz respeito à produção e sobrevivência de mudas em gerações subsequentes. Para compreender os impactos de longo prazo da exploração madeireira e da restauração, acompanhamos a diversidade, sobrevivência e características de plântulas que germinaram imediatamente após uma frutificação em massa no norte de Bornéu, em florestas com e sem exploração de madeira, 30-35 anos após o fim da extração. Monitoramos 5119 mudas desde a germinação por aproximadamente 1,5 anos em uma paisagem mista de florestas não exploradas (UL), florestas exploradas em regeneração natural (NR) e florestas exploradas restauradas ativamente por meio de tratamentos silviculturais de reabilitação (AR), 15-27 anos após a restauração. Medimos 14 traços funcionais de folhas, raízes e alocação de biomassa em 399 mudas de 15 espécies. Logo após a frutificação, as florestas UL e AR apresentaram densidades de mudas mais altas do que as florestas NR, mas a sobrevivência foi mais baixa nas florestas AR nos primeiros seis meses. A composição da comunidade diferiu entre os tipos de floresta; as florestas AR e NR teviram menor riqueza de espécies e menor equidade do que as florestas UL 5-6 meses após a frutificação, mas não diferiram entre si. As diferenças na composição da comunidade alteraram os valores de média ponderada pela comunidade das características entre os tipos de floresta com maior alocação de biomassa radicular nas florestas NR em relação às florestas UL. As características influenciaram a mortalidade aproximadamente 3 meses após a frutificação, com traços mais aquisitivos maior investimento em biomassa relativa acima do solo nas florestas AR em relação às florestas UL. Nossas descobertas de redução na sobrevivência e diversidade de plântulas sugerem que há longos retardos no recrutamento após o fim da exploração de madeira, particularmente para alguns táxons. A restauração ativa de florestas exploradas recupera a produção inicial de plântulas, mas a mortalidade elevada nas florestas AR diminui a eficácia da restauração ativa no melhorio do recrutamento e da diversidade das comunidades de mudas. Isso sugere que as práticas atuais de restauração ativa podem não superar as barreiras à regeneração em florestas exploradas, o que pode levar a mudanças de longo prazo nas comunidades florestais no futuro.

Divergence of hydraulic traits among tropical forest trees across topographic and vertical environment gradients in Borneo.

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.

Incorporating connectivity into conservation planning for the optimal representation of multiple species and ecosystem services.

Conservation planning tends to focus on protecting species' ranges or landscape connectivity but seldom both-particularly in the case of diverse taxonomic assemblages and multiple planning goals. Therefore, information on potential trade-offs between maintaining landscape connectivity and achieving other conservation objectives is lacking. We developed an optimization approach to prioritize the maximal protection of species' ranges, ecosystem types, and forest carbon stocks, while also including habitat connectivity for range-shifting species and dispersal corridors to link protected area. We applied our approach to Sabah, Malaysia, where the state government mandated an increase in protected-area coverage of approximately 305,000 ha but did not specify where new protected areas should be. Compared with a conservation planning approach that did not incorporate the 2 connectivity features, our approach increased the protection of dispersal corridors and elevational connectivity by 13% and 21%, respectively. Coverage of vertebrate and plant species' ranges and forest types were the same whether connectivity was included or excluded. Our approach protected 2% less forest carbon and 3% less butterfly range than when connectivity features were not included. Hence, the inclusion of connectivity into conservation planning can generate large increases in the protection of landscape connectivity with minimal loss of representation of other conservation targets.

Incorporación de la Conectividad a la Planeación de la Conservación para la Representación Óptima de Especies Múltiples y Servicios Ambientales Resumen Las tendencias de planeación de la conservación tienden a enfocarse en la protección de la distribución geográfica de las especies o en la conectividad de paisajes, pero rara vez se enfocan en ambas - particularmente para el caso de los ensamblajes taxonómicos y las metas múltiples de planeación. Por lo tanto, hay carencias en la información sobre las compensaciones potenciales entre mantener la conectividad de los paisajes y alcanzar otros objetivos de conservación. Desarrollamos una estrategia de optimización para priorizar la protección máxima de la distribución de las especies, los tipos de ecosistemas y los stocks de carbono de los bosques, a la vez que incluimos la conectividad del hábitat para las especies que modifican su distribución y los corredores de dispersión para conectar el área protegida. Aplicamos nuestra estrategia en Sabah, Malasia, en donde el gobierno estatal ordenó un incremento de ∼305, 000 ha en la cobertura de áreas protegidas sin especificar la ubicación de las nuevas áreas protegidas. En comparación con una estrategia de planeación de la conservación que no incorporó las dos características de la conectividad, nuestra estrategia incrementó la protección de los corredores de dispersión y la conectividad altitudinal en un 13% y 21% respectivamente. La cobertura de la distribución de las especies de plantas y vertebrados y de los tipos de bosque fue la misma con o sin la inclusión de la conectividad. Nuestra estrategia protegió 2% menos del carbono forestal y 3% menos de la distribución de mariposas que cuando no se incluyeron las características de conectividad en la estrategia. Por lo tanto, incluir a la conectividad en la planeación de la conservación puede generar grandes incrementos en la protección de la conectividad del paisaje con una pérdida mínima de representación para los demás objetivos de conservación.

Reconciling the contribution of environmental and stochastic structuring of tropical forest diversity through the lens of imaging spectroscopy.

Both niche and stochastic dispersal processes structure the extraordinary diversity of tropical plants, but determining their relative contributions has proven challenging. We address this question using airborne imaging spectroscopy to estimate canopy ß-diversity for an extensive region of a Bornean rainforest and challenge these data with models incorporating niches and dispersal. We show that remotely sensed and field-derived estimates of pairwise dissimilarity in community composition are closely matched, proving the applicability of imaging spectroscopy to provide ß-diversity data for entire landscapes of over 1000 ha containing contrasting forest types. Our model reproduces the empirical data well and shows that the ecological processes maintaining tropical forest diversity are scale dependent. Patterns of ß-diversity are shaped by stochastic dispersal processes acting locally whilst environmental processes act over a wider range of scales.

Topography shapes the structure, composition and function of tropical forest landscapes.

Topography is a key driver of tropical forest structure and composition, as it constrains local nutrient and hydraulic conditions within which trees grow. Yet, we do not fully understand how changes in forest physiognomy driven by topography impact other emergent properties of forests, such as their aboveground carbon density (ACD). Working in Borneo - at a site where 70-m-tall forests in alluvial valleys rapidly transition to stunted heath forests on nutrient-depleted dip slopes - we combined field data with airborne laser scanning and hyperspectral imaging to characterise how topography shapes the vertical structure, wood density, diversity and ACD of nearly 15 km2 of old-growth forest. We found that subtle differences in elevation - which control soil chemistry and hydrology - profoundly influenced the structure, composition and diversity of the canopy. Capturing these processes was critical to explaining landscape-scale heterogeneity in ACD, highlighting how emerging remote sensing technologies can provide new insights into long-standing ecological questions.

The value of biodiversity for the functioning of tropical forests: insurance effects during the first decade of the Sabah biodiversity experiment.

One of the main environmental threats in the tropics is selective logging, which has degraded large areas of forest. In southeast Asia, enrichment planting with seedlings of the dominant group of dipterocarp tree species aims to accelerate restoration of forest structure and functioning. The role of tree diversity in forest restoration is still unclear, but the 'insurance hypothesis' predicts that in temporally and spatially varying environments planting mixtures may stabilize functioning owing to differences in species traits and ecologies. To test for potential insurance effects, we analyse the patterns of seedling mortality and growth in monoculture and mixture plots over the first decade of the Sabah biodiversity experiment. Our results reveal the species differences required for potential insurance effects including a trade-off in which species with denser wood have lower growth rates but higher survival. This trade-off was consistent over time during the first decade, but growth and mortality varied spatially across our 500 ha experiment with species responding to changing conditions in different ways. Overall, average survival rates were extreme in monocultures than mixtures consistent with a potential insurance effect in which monocultures of poorly surviving species risk recruitment failure, whereas monocultures of species with high survival have rates of self-thinning that are potentially wasteful when seedling stocks are limited. Longer-term monitoring as species interactions strengthen will be needed to more comprehensively test to what degree mixtures of species spread risk and use limited seedling stocks more efficiently to increase diversity and restore ecosystem structure and functioning.

The relationship between leaf area index and microclimate in tropical forest and oil palm plantation: Forest disturbance drives changes in microclimate.

Land use change is a major threat to biodiversity. One mechanism by which land use change influences biodiversity and ecological processes is through changes in the local climate. Here, the relationships between leaf area index and five climate variables - air temperature, relative humidity, vapour pressure deficit, specific humidity and soil temperature - are investigated across a range of land use types in Borneo, including primary tropical forest, logged forest and oil palm plantation. Strong correlations with the leaf area index are found for the mean daily maximum air and soil temperatures, the mean daily maximum vapour pressure deficit and the mean daily minimum relative humidity. Air beneath canopies with high leaf area index is cooler and has higher relative humidity during the day. Forest microclimate is also found to be less variable for sites with higher leaf area indices. Primary forest is found to be up to 2.5 °C cooler than logged forest and up to 6.5 °C cooler than oil palm plantations. Our results indicate that leaf area index is a useful parameter for predicting the effects of vegetation upon microclimate, which could be used to make small scale climate predictions based on remotely sensed data.

Soils on exposed Sunda shelf shaped biogeographic patterns in the equatorial forests of Southeast Asia.

The marked biogeographic difference between western (Malay Peninsula and Sumatra) and eastern (Borneo) Sundaland is surprising given the long time that these areas have formed a single landmass. A dispersal barrier in the form of a dry savanna corridor during glacial maxima has been proposed to explain this disparity. However, the short duration of these dry savanna conditions make it an unlikely sole cause for the biogeographic pattern. An additional explanation might be related to the coarse sandy soils of central Sundaland. To test these two nonexclusive hypotheses, we performed a floristic cluster analysis based on 111 tree inventories from Peninsular Malaysia, Sumatra, and Borneo. We then identified the indicator genera for clusters that crossed the central Sundaland biogeographic boundary and those that did not cross and tested whether drought and coarse-soil tolerance of the indicator genera differed between them. We found 11 terminal floristic clusters, 10 occurring in Borneo, 5 in Sumatra, and 3 in Peninsular Malaysia. Indicator taxa of clusters that occurred across Sundaland had significantly higher coarse-soil tolerance than did those from clusters that occurred east or west of central Sundaland. For drought tolerance, no such pattern was detected. These results strongly suggest that exposed sandy sea-bed soils acted as a dispersal barrier in central Sundaland. However, we could not confirm the presence of a savanna corridor. This finding makes it clear that proposed biogeographic explanations for plant and animal distributions within Sundaland, including possible migration routes for early humans, need to be reevaluated.

Tropical tree ectomycorrhiza are distributed independently of soil nutrients.

Mycorrhizae, a form of plant-fungal symbioses, mediate vegetation impacts on ecosystem functioning. Climatic effects on decomposition and soil quality are suggested to drive mycorrhizal distributions, with arbuscular mycorrhizal plants prevailing in low-latitude/high-soil-quality areas and ectomycorrhizal (EcM) plants in high-latitude/low-soil-quality areas. However, these generalizations, based on coarse-resolution data, obscure finer-scale variations and result in high uncertainties in the predicted distributions of mycorrhizal types and their drivers. Using data from 31 lowland tropical forests, both at a coarse scale (mean-plot-level data) and fine scale (20 × 20 metres from a subset of 16 sites), we demonstrate that the distribution and abundance of EcM-associated trees are independent of soil quality. Resource exchange differences among mycorrhizal partners, stemming from diverse evolutionary origins of mycorrhizal fungi, may decouple soil fertility from the advantage provided by mycorrhizal associations. Additionally, distinct historical biogeographies and diversification patterns have led to differences in forest composition and nutrient-acquisition strategies across three major tropical regions. Notably, Africa and Asia's lowland tropical forests have abundant EcM trees, whereas they are relatively scarce in lowland neotropical forests. A greater understanding of the functional biology of mycorrhizal symbiosis is required, especially in the lowland tropics, to overcome biases from assuming similarity to temperate and boreal regions.

A Demi-Decade of Mammal Research: A Rapid Assessment within the Heart of Borneo in Sabah.

Sabah contributes 4.2 million hectares to the total Heart of Borneo (HoB) areas. Some of the forest reserves in the HoB are newly gazetted as Totally Protected Forest. Hence, their mammal diversity has to be comprehensively documented. This study aims to record the presence of terrestrial mammal species, and assess the prevalence of poaching in selected forest reserves within the Sabah HoB area. A total of 15 forest reserves were surveyed within a 5-year timeframe which recorded 60 terrestrial mammal species, including 21 Bornean endemics. The variation in total enumerated mammal species in the study sites may be derived from unequal sampling efforts, geographical factors and anthropogenic influences. The intensity of poaching within the study sites is high. Though this study is a rapid assessment, it created baseline information for mammal diversity in some of the least studied forest reserves in Sabah, important for conservation of its terrestrial mammals.

Sabah menyumbang 4.2 juta hektar kepada keluasan kawasan Heart of Borneo (HoB). Beberapa hutan simpan dalam kawasan HoB baru sahaja diwartakan sebagai Kawasan Terlindung Sepenuhnya. Justeru, kepelbagaian mamalia dalam kawasan hutan simpan tersebut perlu didokumentasi secara menyeluruh. Kajian ini bertujuan merekod kewujudan mamalia daratan dan menilai kelaziman aktiviti pemburuan haram di hutan simpan terpilih dalam kawasan HoB di Sabah. Sejumlah 15 hutan simpan telah ditinjau dalam tempoh masa lima tahun. Sebanyak 60 spesies mamalia daratan termasuk 21 spesies endemik di Borneo telah direkodkan. Perbezaaan jumlah spesies mamalia yang direkod di hutan simpan terpilih mungkin disebabkan oleh ketidaksamaan usaha pensampelan, faktor geografi dan tekanan antropogenik. Intensiti pemburuan haram dalam kawasan kajian adalah tinggi. Walaupun kajian ini hanyalah tinjauan pantas, namun ia menghasilkan maklumat asas kepelbagaian mamalia dalam hutan simpan yang kurang diterokai serta menunjukkan kebergunaan pendekatan tinjauan pantas dalam memulihara mamalia daratan.

Global patterns and climatic controls of forest structural complexity.

The complexity of forest structures plays a crucial role in regulating forest ecosystem functions and strongly influences biodiversity. Yet, knowledge of the global patterns and determinants of forest structural complexity remains scarce. Using a stand structural complexity index based on terrestrial laser scanning, we quantify the structural complexity of boreal, temperate, subtropical and tropical primary forests. We find that the global variation of forest structural complexity is largely explained by annual precipitation and precipitation seasonality (R² = 0.89). Using the structural complexity of primary forests as benchmark, we model the potential structural complexity across biomes and present a global map of the potential structural complexity of the earth´s forest ecoregions. Our analyses reveal distinct latitudinal patterns of forest structure and show that hotspots of high structural complexity coincide with hotspots of plant diversity. Considering the mechanistic underpinnings of forest structural complexity, our results suggest spatially contrasting changes of forest structure with climate change within and across biomes.

Herbivory of tropical rain forest tree seedlings correlates with future mortality.

Tree seedlings in tropical rain forests are subject to both damage from natural enemies and intense interspecific competition. This leads to a trade-off in investment between defense and growth, and it is likely that tree species specialized to particular habitats tailor this balance to correspond with local resource availability. It has also been suggested that differential herbivore impacts among tree species may drive habitat segregation, favoring species adapted to particular resource conditions. In order to test these predictions, a reciprocal transplant experiment in Sabah, Malaysia, was established with seedlings of five species of Dipterocarpaceae. These were specialized to either alluvial (Hopea nervosa, Parashorea tomentella) or sandstone soils (Shorea multiflora, H. beccariana), or were locally absent (S. fallax). A total of 3000 seedlings were planted in paired gap and understory plots in five sites on alluvial and sandstone soils. Half of all seedlings were fertilized. Seedling growth and mortality were recorded in regular samples over 3.5 years, and rates of insect herbivore damage were estimated from censuses of foliar tissue loss on marked mature leaves and available young leaves. Greater herbivory rates on mature leaves had no measurable effects on seedling growth but were associated with a significantly increased likelihood of mortality during the following year. In contrast, new-leaf herbivory rates correlated with neither growth nor mortality. There were no indications of differential impacts of herbivory among the five species, nor between experimental treatments. Herbivory was not shown to influence segregation of species between soil types, although it may contribute toward differential survival among light habitats. Natural rates of damage were substantially lower than have been shown to influence tree seedling growth and mortality in previous manipulative studies.

The value of rehabilitating logged rainforest for birds.

The recent advent of carbon crediting has led to a rapid rise in biosequestration projects that seek to remove carbon from the atmosphere through afforestation and forest rehabilitation. Such projects also present an important potential opportunity to reverse biodiversity losses resulting from deforestation and forest degradation, but the biodiversity benefits of different forms of biosequestration have not been considered adequately. We captured birds in mist nets to examine the effects of rehabilitation of logged forest on birds in Sabah, Borneo, and to test the hypothesis that rehabilitation restores avian assemblages within regenerating forest to a condition closer to that seen in unlogged forest. Species richness and diversity were similar in unlogged and rehabilitated forest, but significantly lower in naturally regenerating forest. Rehabilitation resulted in a relatively rapid recovery of populations of insectivores within logged forest, especially those species that forage by sallying, but had a marked adverse effect on frugivores and possibly reduced the overall abundance of birds within regenerating forest. In view of these results, we advocate increased management for heterogeneity within rehabilitated forests, but we strongly urge an increased role for forest rehabilitation in the design and implementation of a biodiversity-friendly carbon-offsetting market.

Author Correction: Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edges.

The original version of this Article contained an error in the third sentence of the abstract and incorrectly read "Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 year-1 (95% CI 0.14-0.72, mean period 1988-2010) above-ground live biomass", rather than the correct "Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 year-1 (95% CI 0.14-0.72, mean period 1988-2010) in above-ground live biomass carbon". This has now been corrected in both the PDF and HTML versions of the Article.

Long-term carbon sink in Borneo's forests halted by drought and vulnerable to edge effects.

Less than half of anthropogenic carbon dioxide emissions remain in the atmosphere. While carbon balance models imply large carbon uptake in tropical forests, direct on-the-ground observations are still lacking in Southeast Asia. Here, using long-term plot monitoring records of up to half a century, we find that intact forests in Borneo gained 0.43 Mg C ha-1 per year (95% CI 0.14-0.72, mean period 1988-2010) above-ground live biomass. These results closely match those from African and Amazonian plot networks, suggesting that the world's remaining intact tropical forests are now en masse out-of-equilibrium. Although both pan-tropical and long-term, the sink in remaining intact forests appears vulnerable to climate and land use changes. Across Borneo the 1997-1998 El Niño drought temporarily halted the carbon sink by increasing tree mortality, while fragmentation persistently offset the sink and turned many edge-affected forests into a carbon source to the atmosphere.

Logging cuts the functional importance of invertebrates in tropical rainforest.

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests.

A trait-based trade-off between growth and mortality: evidence from 15 tropical tree species using size-specific relative growth rates.

A life-history trade-off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged-over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species-specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed-effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed-effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade-off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade-off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.