Ghost roads and the destruction of Asia-Pacific tropical forests.

Roads are expanding at the fastest pace in human history. This is the case especially in biodiversity-rich tropical nations, where roads can result in forest loss and fragmentation, wildfires, illicit land invasions and negative societal effects1-5. Many roads are being constructed illegally or informally and do not appear on any existing road map6-10; the toll of such 'ghost roads' on ecosystems is poorly understood. Here we use around 7,000 h of effort by trained volunteers to map ghost roads across the tropical Asia-Pacific region, sampling 1.42 million plots, each 1 km2 in area. Our intensive sampling revealed a total of 1.37 million km of roads in our plots-from 3.0 to 6.6 times more roads than were found in leading datasets of roads globally. Across our study area, road building almost always preceded local forest loss, and road density was by far the strongest correlate11 of deforestation out of 38 potential biophysical and socioeconomic covariates. The relationship between road density and forest loss was nonlinear, with deforestation peaking soon after roads penetrate a landscape and then declining as roads multiply and remaining accessible forests largely disappear. Notably, after controlling for lower road density inside protected areas, we found that protected areas had only modest additional effects on preventing forest loss, implying that their most vital conservation function is limiting roads and road-related environmental disruption. Collectively, our findings suggest that burgeoning, poorly studied ghost roads are among the gravest of all direct threats to tropical forests.

Global dominance of lianas over trees is driven by forest disturbance, climate and topography.

Growing evidence suggests that liana competition with trees is threatening the global carbon sink by slowing the recovery of forests following disturbance. A recent theory based on local and regional evidence further proposes that the competitive success of lianas over trees is driven by interactions between forest disturbance and climate. We present the first global assessment of liana-tree relative performance in response to forest disturbance and climate drivers. Using an unprecedented dataset, we analysed 651 vegetation samples representing 26,538 lianas and 82,802 trees from 556 unique locations worldwide, derived from 83 publications. Results show that lianas perform better relative to trees (increasing liana-to-tree ratio) when forests are disturbed, under warmer temperatures and lower precipitation and towards the tropical lowlands. We also found that lianas can be a critical factor hindering forest recovery in disturbed forests experiencing liana-favourable climates, as chronosequence data show that high competitive success of lianas over trees can persist for decades following disturbances, especially when the annual mean temperature exceeds 27.8°C, precipitation is less than 1614 mm and climatic water deficit is more than 829 mm. These findings reveal that degraded tropical forests with environmental conditions favouring lianas are disproportionately more vulnerable to liana dominance and thus can potentially stall succession, with important implications for the global carbon sink, and hence should be the highest priority to consider for restoration management.

Des preuves de plus en plus nombreuses suggèrent que la competition entre lianes et les arbres menace le puits de carbone mondial en ralentissant la récupération des forêts après une perturbation. Une théorie récente, fondée sur des observations locales et régionales, propose en outre que le succès compétitif des lianes sur les arbres est dû aux interactions entre la perturbation forestière et le climat. Nous présentons la première évaluation mondiale de la performance relative des lianes par rapport aux arbres en réponse aux perturbations forestières et aux facteurs climatiques. En utilisant un ensemble de données sans précédent, nous avons analysé 651 échantillons de végétation représentant 26,538 lianes et 82,802 arbres, issus de 556 emplacements uniques dans le monde entier, tirés de 83 publications. Les résultats montrent que les lianes ont de meilleure performances par rapport aux arbres (augmentation du ratio liane-arbre) lorsque les forêts sont perturbées, sous des zones chaudes aves précipitations faibles, et vers les basses altitudes tropicales. Nous avons également constaté que les lianes peuvent être un facteur critique entravant la récupération des forêts dans les forêts perturbées connaissant des climats favorables aux lianes, car les données de chronoséquence montrent que le succès compétitif élevé des lianes sur les arbres peut persister pendant des décennies après les perturbations, surtout lorsque la température annuelle moyenne dépasse 27.8°C, que les précipitations sont inférieures à 1614 mm et que le déficit hydrique climatique est supérieur à 829 mm. Ces découvertes révèlent que les forêts tropicales dégradées avec des conditions environnementales favorables aux lianes sont disproportionnellement plus vulnérables à la dominance des lianes, et peuvent ainsi potentiellement entraver la succession, avec d'importantes implications pour le puits de carbone mondial et devraient donc être la plus haute priorité à considérer pour la gestion de la restauration.

A global meta-analysis of the impacts of forest fragmentation on biotic mutualisms and antagonisms.

Forest fragmentation is a grave threat to biodiversity. Forests are becoming increasingly fragmented with more than 70% now < 1 km from forest edge. Although much is known about the effects of forest fragmentation on individual species, much less is understood about its effects on species interactions (i.e., mutualisms, antagonisms, etc.). In 2014, a previous meta-analysis assessed the impacts of forest fragmentation on different species interactions, across 82 studies. We pooled the previous data with data published in the last 10 years (combined total 104 studies and 168 effect sizes). We compared the new set of publications (22 studies and 32 effect sizes) with the old set to evaluate potential changes in species interactions over time given the global increase in fragmentation rates. Mutualisms were more negatively affected by forest fragmentation than antagonisms (p < 0.0001). Edge effects, fragment size, and degradation negatively affected mutualisms, but not antagonisms, a different finding from the original meta-analysis. Parasitic interactions increased as fragment size decreased (p < 0.0001)-an intriguing result at variance with earlier studies. New publications showed a more negative mean effect size of forest fragmentation on mutualisms than old publications. Although research is still limited for some interactions, we identified an important scientific trend: current research tends to focus on antagonisms. We concluded that forest fragmentation disrupts important species interactions and that this disruption has increased over time.

Metaanálisis Mundial del Impacto de la Fragmentación de Bosques sobre el Mutualismo y Antagonismo Biótico Resumen La fragmentación del bosque es una amenaza grave para la biodiversidad. Los bosques están más fragmentados, pues más del 70% tienen < 1 km a partir del borde del bosque. Aunque hay mucha información del efecto de la fragmentación sobre las especies, hay poco conocimiento de sus efectos sobre las interacciones entre especies (mutualismo, antagonismo, etc.). Un metaanálisis realizado en 2014 por evaluó en 82 estudios el impacto de la fragmentación del bosque sobre las diferentes interacciones entre especies. Juntamos estos datos con datos publicados en los últimos diez años (total combinado de 104 estudios y 168 tamaños de efecto). Comparamos el grupo nuevo de publicaciones (22 estudios y 32 tamaños de efecto) con el grupo de para evaluar los cambios potenciales en las interacciones entre especies con el tiempo dado el incremento mundial de la tasa de fragmentación. El mutualismo fue más afectado negativamente por la fragmentación del bosque que los antagonismos (p < 0.0001). El efecto del borde, tamaño del fragmento y la degradación tuvieron un efecto negativo sobre el mutualismo, pero no sobre el antagonismo, un resultado diferente al del metaanálisis original. Las interacciones parasitarias incrementaron conforme se redujo el tamaño del fragmento (p < 0.0001)-un resultado intrigante en discrepancia con los primeros resultados. Las publicaciones recientes mostraron un tamaño promedio de efecto de la fragmentación del bosque más negativo para el mutualismo que las publicaciones antiguas. Aunque hay poca investigación sobre algunas interacciones, identificamos una tendencia científica importante: la investigación actual tiende a enfocarse en los antagonismos. Concluimos que la fragmentación del bosque altera las interacciones importantes entre especies y que este cambio ha aumentado con el tiempo.

A global strategy for road building.

The number and extent of roads will expand dramatically this century. Globally, at least 25 million kilometres of new roads are anticipated by 2050; a 60% increase in the total length of roads over that in 2010. Nine-tenths of all road construction is expected to occur in developing nations, including many regions that sustain exceptional biodiversity and vital ecosystem services. Roads penetrating into wilderness or frontier areas are a major proximate driver of habitat loss and fragmentation, wildfires, overhunting and other environmental degradation, often with irreversible impacts on ecosystems. Unfortunately, much road proliferation is chaotic or poorly planned, and the rate of expansion is so great that it often overwhelms the capacity of environmental planners and managers. Here we present a global scheme for prioritizing road building. This large-scale zoning plan seeks to limit the environmental costs of road expansion while maximizing its benefits for human development, by helping to increase agricultural production, which is an urgent priority given that global food demand could double by mid-century. Our analysis identifies areas with high environmental values where future road building should be avoided if possible, areas where strategic road improvements could promote agricultural development with relatively modest environmental costs, and 'conflict areas' where road building could have sizeable benefits for agriculture but with serious environmental damage. Our plan provides a template for proactively zoning and prioritizing roads during the most explosive era of road expansion in human history.

Persistent effects of fragmentation on tropical rainforest canopy structure after 20 yr of isolation.

Assessing the persistent impacts of fragmentation on aboveground structure of tropical forests is essential to understanding the consequences of land use change for carbon storage and other ecosystem functions. We investigated the influence of edge distance and fragment size on canopy structure, aboveground woody biomass (AGB), and AGB turnover in the Biological Dynamics of Forest Fragments Project (BDFFP) in central Amazon, Brazil, after 22+ yr of fragment isolation, by combining canopy variables collected with portable canopy profiling lidar and airborne laser scanning surveys with long-term forest inventories. Forest height decreased by 30% at edges of large fragments (>10 ha) and interiors of small fragments (<3 ha). In larger fragments, canopy height was reduced up to 40 m from edges. Leaf area density profiles differed near edges: the density of understory vegetation was higher and midstory vegetation lower, consistent with canopy reorganization via increased regeneration of pioneers following post-fragmentation mortality of large trees. However, canopy openness and leaf area index remained similar to control plots throughout fragments, while canopy spatial heterogeneity was generally lower at edges. AGB stocks and fluxes were positively related to canopy height and negatively related to spatial heterogeneity. Other forest structure variables typically used to assess the ecological impacts of fragmentation (basal area, density of individuals, and density of pioneer trees) were also related to lidar-derived canopy surface variables. Canopy reorganization through the replacement of edge-sensitive species by disturbance-tolerant ones may have mitigated the biomass loss effects due to fragmentation observed in the earlier years of BDFFP. Lidar technology offered novel insights and observational scales for analysis of the ecological impacts of fragmentation on forest structure and function, specifically aboveground biomass storage.

Averting biodiversity collapse in tropical forest protected areas.

The rapid disruption of tropical forests probably imperils global biodiversity more than any other contemporary phenomenon. With deforestation advancing quickly, protected areas are increasingly becoming final refuges for threatened species and natural ecosystem processes. However, many protected areas in the tropics are themselves vulnerable to human encroachment and other environmental stresses. As pressures mount, it is vital to know whether existing reserves can sustain their biodiversity. A critical constraint in addressing this question has been that data describing a broad array of biodiversity groups have been unavailable for a sufficiently large and representative sample of reserves. Here we present a uniquely comprehensive data set on changes over the past 20 to 30 years in 31 functional groups of species and 21 potential drivers of environmental change, for 60 protected areas stratified across the world's major tropical regions. Our analysis reveals great variation in reserve 'health': about half of all reserves have been effective or performed passably, but the rest are experiencing an erosion of biodiversity that is often alarmingly widespread taxonomically and functionally. Habitat disruption, hunting and forest-product exploitation were the strongest predictors of declining reserve health. Crucially, environmental changes immediately outside reserves seemed nearly as important as those inside in determining their ecological fate, with changes inside reserves strongly mirroring those occurring around them. These findings suggest that tropical protected areas are often intimately linked ecologically to their surrounding habitats, and that a failure to stem broad-scale loss and degradation of such habitats could sharply increase the likelihood of serious biodiversity declines.

Brazil's worst mining disaster: Corporations must be compelled to pay the actual environmental costs.

In November 2015, a large mine-tailing dam owned by Samarco Corporation collapsed in Brazil, generating a massive wave of toxic mud that spread down the Doce River, killing 20 people and affecting biodiversity across hundreds of kilometers of river, riparian lands, and Atlantic coast. Besides the disaster's serious human and socioeconomic tolls, we estimate the regional loss of environmental services to be ~US$521 million per year. Although our estimate is conservative, it is still six times higher than the fine imposed on Samarco by Brazilian environmental authorities. To reduce such disparities between estimated damages and levied fines, we advocate for an environmental bond policy that considers potential risks and environmental services that could possibly be impacted by irresponsible mining activity. Environmental bonds and insurance are commonly used policy instruments in many countries, but there are no clear environmental bond policies in Brazil. Environmental bonds are likely to be more effective at securing environmental restitution than post-disaster fines, which generally are inadequate and often unpaid. We estimate that at least 126 mining dams in Brazil are vulnerable to failure in the forthcoming years. Any such event could have severe social-environmental consequences, underscoring the need for effective disaster-management strategies for large-scale mining operations.

Primary forests are irreplaceable for sustaining tropical biodiversity.

Human-driven land-use changes increasingly threaten biodiversity, particularly in tropical forests where both species diversity and human pressures on natural environments are high. The rapid conversion of tropical forests for agriculture, timber production and other uses has generated vast, human-dominated landscapes with potentially dire consequences for tropical biodiversity. Today, few truly undisturbed tropical forests exist, whereas those degraded by repeated logging and fires, as well as secondary and plantation forests, are rapidly expanding. Here we provide a global assessment of the impact of disturbance and land conversion on biodiversity in tropical forests using a meta-analysis of 138 studies. We analysed 2,220 pairwise comparisons of biodiversity values in primary forests (with little or no human disturbance) and disturbed forests. We found that biodiversity values were substantially lower in degraded forests, but that this varied considerably by geographic region, taxonomic group, ecological metric and disturbance type. Even after partly accounting for confounding colonization and succession effects due to the composition of surrounding habitats, isolation and time since disturbance, we find that most forms of forest degradation have an overwhelmingly detrimental effect on tropical biodiversity. Our results clearly indicate that when it comes to maintaining tropical biodiversity, there is no substitute for primary forests.

Selective logging in tropical forests decreases the robustness of liana-tree interaction networks to the loss of host tree species.

Selective logging is one of the major drivers of tropical forest degradation, causing important shifts in species composition. Whether such changes modify interactions between species and the networks in which they are embedded remain fundamental questions to assess the 'health' and ecosystem functionality of logged forests. We focus on interactions between lianas and their tree hosts within primary and selectively logged forests in the biodiversity hotspot of Malaysian Borneo. We found that lianas were more abundant, had higher species richness, and different species compositions in logged than in primary forests. Logged forests showed heavier liana loads disparately affecting slow-growing tree species, which could exacerbate the loss of timber value and carbon storage already associated with logging. Moreover, simulation scenarios of host tree local species loss indicated that logging might decrease the robustness of liana-tree interaction networks if heavily infested trees (i.e. the most connected ones) were more likely to disappear. This effect is partially mitigated in the short term by the colonization of host trees by a greater diversity of liana species within logged forests, yet this might not compensate for the loss of preferred tree hosts in the long term. As a consequence, species interaction networks may show a lagged response to disturbance, which may trigger sudden collapses in species richness and ecosystem function in response to additional disturbances, representing a new type of 'extinction debt'.

Evolutionary heritage influences Amazon tree ecology.

Lineages tend to retain ecological characteristics of their ancestors through time. However, for some traits, selection during evolutionary history may have also played a role in determining trait values. To address the relative importance of these processes requires large-scale quantification of traits and evolutionary relationships among species. The Amazonian tree flora comprises a high diversity of angiosperm lineages and species with widely differing life-history characteristics, providing an excellent system to investigate the combined influences of evolutionary heritage and selection in determining trait variation. We used trait data related to the major axes of life-history variation among tropical trees (e.g. growth and mortality rates) from 577 inventory plots in closed-canopy forest, mapped onto a phylogenetic hypothesis spanning more than 300 genera including all major angiosperm clades to test for evolutionary constraints on traits. We found significant phylogenetic signal (PS) for all traits, consistent with evolutionarily related genera having more similar characteristics than expected by chance. Although there is also evidence for repeated evolution of pioneer and shade tolerant life-history strategies within independent lineages, the existence of significant PS allows clearer predictions of the links between evolutionary diversity, ecosystem function and the response of tropical forests to global change.

Variation in stem mortality rates determines patterns of above-ground biomass in Amazonian forests: implications for dynamic global vegetation models.

Understanding the processes that determine above-ground biomass (AGB) in Amazonian forests is important for predicting the sensitivity of these ecosystems to environmental change and for designing and evaluating dynamic global vegetation models (DGVMs). AGB is determined by inputs from woody productivity [woody net primary productivity (NPP)] and the rate at which carbon is lost through tree mortality. Here, we test whether two direct metrics of tree mortality (the absolute rate of woody biomass loss and the rate of stem mortality) and/or woody NPP, control variation in AGB among 167 plots in intact forest across Amazonia. We then compare these relationships and the observed variation in AGB and woody NPP with the predictions of four DGVMs. The observations show that stem mortality rates, rather than absolute rates of woody biomass loss, are the most important predictor of AGB, which is consistent with the importance of stand size structure for determining spatial variation in AGB. The relationship between stem mortality rates and AGB varies among different regions of Amazonia, indicating that variation in wood density and height/diameter relationships also influences AGB. In contrast to previous findings, we find that woody NPP is not correlated with stem mortality rates and is weakly positively correlated with AGB. Across the four models, basin-wide average AGB is similar to the mean of the observations. However, the models consistently overestimate woody NPP and poorly represent the spatial patterns of both AGB and woody NPP estimated using plot data. In marked contrast to the observations, DGVMs typically show strong positive relationships between woody NPP and AGB. Resolving these differences will require incorporating forest size structure, mechanistic models of stem mortality and variation in functional composition in DGVMs.

Would protecting tropical forest fragments provide carbon and biodiversity cobenefits under REDD+?

Tropical forests store vast amounts of carbon and are the most biodiverse terrestrial habitats, yet they are being converted and degraded at alarming rates. Given global shortfalls in the budgets required to prevent carbon and biodiversity loss, we need to seek solutions that simultaneously address both issues. Of particular interest are carbon-based payments under the Reducing Emissions from Deforestation and Forest Degradation (REDD+) mechanism to also conserve biodiversity at no additional cost. One potential is for REDD+ to protect forest fragments, especially within biomes where contiguous forest cover has diminished dramatically, but we require empirical tests of the strength of any carbon and biodiversity cobenefits in such fragmented systems. Using the globally threatened Atlantic Forest landscape, we measured above-ground carbon stocks within forest fragments spanning 13 to 23 442 ha in area and with different degrees of isolation. We related these stocks to tree community structure and to the richness and abundance of endemic and IUCN Red-listed species. We found that increasing fragment size has a positive relationship with above-ground carbon stock and with abundance of IUCN Red-listed species and tree community structure. We also found negative relationships between distance from large forest block and tree community structure, endemic species richness and abundance, and IUCN Red-listed species abundance. These resulted in positive congruence between carbon stocks and Red-listed species, and the abundance and richness of endemic species, demonstrating vital cobenefits. As such, protecting forest fragments in hotspots of biodiversity, particularly larger fragments and those closest to sources, offers important carbon and biodiversity cobenefits. More generally, our results suggest that macroscale models of cobenefits under REDD+ have likely overlooked key benefits at small scales, indicating the necessity to apply models that include finer-grained assessments in fragmented landscapes rather than using averaged coarse-grained cells.

Fast demographic traits promote high diversification rates of Amazonian trees.

The Amazon rain forest sustains the world's highest tree diversity, but it remains unclear why some clades of trees are hyperdiverse, whereas others are not. Using dated phylogenies, estimates of current species richness and trait and demographic data from a large network of forest plots, we show that fast demographic traits--short turnover times--are associated with high diversification rates across 51 clades of canopy trees. This relationship is robust to assuming that diversification rates are either constant or decline over time, and occurs in a wide range of Neotropical tree lineages. This finding reveals the crucial role of intrinsic, ecological variation among clades for understanding the origin of the remarkable diversity of Amazonian trees and forests.

Long-term changes in liana abundance and forest dynamics in undisturbed Amazonian forests.

Lianas (climbing woody vines) are important structural parasites of tropical trees and may be increasing in abundance in response to global-change drivers. We assessed long-term (-14-year) changes in liana abundance and forest dynamics within 36 1-ha permanent plots spanning -600 km2 of undisturbed rainforest in central Amazonia. Within each plot, we counted each liana stem (> or = 2 cm diameter) and measured its diameter at 1.3 m height, and then used these data to estimate liana aboveground biomass. An initial liana survey was completed in 1997-1999 and then repeated in 2012, using identical methods. Liana abundance in the plots increased by an average of 1.00% +/- 0.88% per year, leading to a highly significant (t = 6.58, df = 35, P < 0.00001) increase in liana stem numbers. Liana biomass rose more slowly over time (0.32% +/- 1.37% per year) and the mean difference between the two sampling intervals was nonsignificant (t = 1.46, df = 35, P = 0.15; paired t tests). Liana size distributions shifted significantly (chi2 = 191, df = 8, P < 0.0001; Chi-square test for independence) between censuses, mainly as a result of a nearly 40% increase in the number of smaller (2-3 cm diameter) lianas, suggesting that lianas recruited rapidly during the study. We used long-term data on rainfall and forest dynamics from our study site to test hypotheses about potential drivers of change in liana communities. Lianas generally increase with rainfall seasonality, but we found no significant trends over time (1997-2012) in five rainfall parameters (total annual rainfall, dry-season rainfall, wet-season rainfall, number of very dry months, CV of monthly rainfall). However, rates of tree mortality and recruitment have increased significantly over time in our plots, and general linear mixed-effect models suggested that lianas were more abundant at sites with higher tree mortality and flatter topography. Rising concentrations of atmospheric CO2, which may stimulate liana growth, might also have promoted liana increases. Our findings clearly support the view that lianas are increasing in abundance in old-growth tropical forests, possibly in response to accelerating forest dynamics and rising CO2 concentrations. The aboveground biomass of trees was lowest in plots with abundant lianas, suggesting that lianas could reduce forest carbon storage and potentially alter forest dynamics if they continue to proliferate.

Land-sharing versus land-sparing logging: reconciling timber extraction with biodiversity conservation.

Selective logging is a major driver of rainforest degradation across the tropics. Two competing logging strategies are proposed to meet timber demands with the least impact on biodiversity: land sharing, which combines timber extraction with biodiversity protection across the concession; and land sparing, in which higher intensity logging is combined with the protection of intact primary forest reserves. We evaluate these strategies by comparing the abundances and species richness of birds, dung beetles and ants in Borneo, using a protocol that allows us to control for both timber yield and net profit across strategies. Within each taxonomic group, more species had higher abundances with land-sparing than land-sharing logging, and this translated into significantly higher species richness within land-sparing concessions. Our results are similar when focusing only on species found in primary forest and restricted in range to Sundaland, and they are independent of the scale of sampling. For each taxonomic group, land-sparing logging was the most promising strategy for maximizing the biological value of logging operations.