Disturbance regime changes the trait distribution,phylogenetic structure and community assembly of tropical rain forests

A rapidly increasing effort to merge functional community ecology and phylogenetic biology has increased our understanding of community assembly. However, studies using both phylogenetic‐ and trait‐based methods have been mainly conducted in old‐growth forests, with fewer studies in human‐disturbed communities, which play an increasingly important role in providing ecosystem services as primary forests are degraded. We used data from 18 1‐ha plots in tropical old‐growth forests and secondary forests with different disturbance histories (logging and shifting cultivation) and vegetation types (tropical lowland and montane forests) on Hainan Island, southern China. The distributions of 11 functional traits were compared among these six forest types. We used a null model approach to assess the effects of disturbance regimes on variation in response and effect traits and community phylogenetic structure across different stem sizes (saplings, treelets, and adult trees) and spatial scales (10–50 m). We found significant differences in the distribution of functional traits in highly disturbed lowland sites versus other forest types. Many individuals in highly disturbed lowland sites were deciduous, spiny, with non‐fleshy fruits and seeds dispersed passively or by wind, and low SLA. The response traits of coexisting species were clustered in all sites except for highly disturbed lowland sites where evenness was evident. There were different distributions of effect traits for saplings and treelets among different forest types but adult trees showed stronger clustering of trait values with increasing spatial scale among all forest types. Phylogenetic clustering predominated across all size classes and spatial scales in the highly disturbed lowland sites, and evenness in other forest types. High disturbance can lead to abiotic filtering, generating a community dominated by closely related species with disturbance‐adapted traits, where biotic interactions play a relatively minor role. In lightly disturbed and old growth forests, multiple processes simultaneously drive the community assembly, but biotic processes dominate at the fine scale.     

Opposing mechanisms affect taxonomic convergence between tree assemblages during tropical forest succession

Whether successional forests converge towards an equilibrium in species composition remains an elusive question, hampered by high idiosyncrasy in successional dynamics. Based on long‐term tree monitoring in second‐growth (SG) and old‐growth (OG) forests in Costa Rica, we show that patterns of convergence between pairs of forest stands depend upon the relative abundance of species exhibiting distinct responses to the successional gradient. For instance, forest generalists contributed to convergence between SG and OG forests, whereas rare species and old‐growth specialists were a source of divergence. Overall, opposing trends in taxonomic similarity among different subsets of species nullified each other, producing a net outcome of stasis over time. Our results offer an explanation for the limited convergence observed between pairwise communities and suggest that rare species and old‐growth specialists may be prone to dispersal limitation, while the dynamics of generalists and second‐growth specialists are more predictable, enhancing resilience in tropical secondary forests.     

Natural regeneration and biodiversity: a global meta‐analysis and implications for spatial planning

Natural regeneration offers a cheaper alternative to active reforestation and has the potential to become the predominant way of restoring degraded tropical landscapes at large‐scale. We conducted a meta‐analysis for tropical regions and quantified the relationships between both ecological and socioeconomic factors and biodiversity responses in naturally regenerating areas. Biogeographic realms, past disturbance, and the human development index (HDI) were used as explanatory variables for biodiversity responses. In addition, we present a case study of large‐scale natural regeneration in the Brazilian Atlantic Forest and identify areas where different forms of restoration would be most suitable. Using our dataset for tropical regions, we showed that natural regeneration was predominantly reported within: the Neotropical realm; areas that were intensively disturbed; and countries with medium HDI. We also found that biodiversity in regenerating forests was more similar to the values found in old growth forests in: countries with either low, high, or very high HDI; less biodiverse realms; and areas of less intensive past disturbance. Our case study from Brazil showed that the level of forest gain resulting from environmental legislation, in particular the Brazilian Forest Code, has been reduced, but remains substantial. Complementary market incentives and financial mechanisms to promote large‐scale natural regeneration in human‐modified agricultural landscapes are also needed. Our analysis provides insights into the factors that promote or limit the recovery of biodiversity in naturally regenerating areas, and aids to identify areas with higher potential for natural regeneration.     

Vegetation survey of the Wasgomuwa National Park,Sri Lanka: analysis of the Wasgomuwa Oya Forest

Three types of forests were recognised (high, disturbed and open) based on the openings in the canopy in a Tropical Monsoonal Forest at the Wasgomuwa National Park, Sri Lanka. The analysis of these forests showed that the species composition varied between forest types. The shrub vegetation, not only was very characteristic but was also a major component in each forest, unlike in the tropical rain forest. The exact role of the shrubs in the dynamics of the tropical monsoon forests is not established, but is suggested that it may act as a buffer during the dry periods under a semi deciduous canopy. The sapling composition differed from the tree vegetation and indicated that the species composition may change with time in this forest. Mosaic theory or the patch dynamics may best explain the dynamics of this tropical monsoon forest which is a mixture of forest types. More studies are required before generalisations can be made of tropical monsoon forests.     

Ice‐storm disturbance and long‐term forest dynamics in the Adirondack Mountains

Ice storms cause periodic disturbance to temperate forests of eastern North America. They are the primary agents of disturbance in some eastern forests. In this paper, a forest gap model is employed to explore consequences of ice storms for the long‐term dynamics of Tsuga canadensis‐northem hardwoods forests. The gap model LINKAGES was modified to simulate periodic ice storm disturbance in the Adirondack Mountains of New York. To adapt the gap model for this purpose, field data on ice storm disturbance are used to develop a polytomous logistic regression model of tree damage. The logistic regression model was then incorporated into the modified forest gap model, LINK ADIR, to determine the type of damage sustained by each simulated tree. The logistic regression model predicts high probabilities of bent boles or severe bole damage (leaning, snapping, or uprooting) in small‐diameter trees, and increasing probability of canopy damage as tree size increases. Canopy damage is most likely on gentle slopes; the probability of severe bole damage increases with increasing slope angle. In the LINKADIR simulations, tree damage type determines the probability of mortality; trees with severe bole damage are assigned the highest mortality rate. LINKADIR predicts Tsuga canadensis dominance in mesophytic old‐growth forests not disturbed by ice storms. When ice storms are simulated, the model predicts Acer saccharum‐dominated forests with higher species richness. These results suggest that ice storms may function as intermediate disturbances that enhance species richness in forested Adirondack landscapes.     

Land snail diversity in post extraction secondary forest reserves in Edo State,Nigeria

Knowledge of the fauna of tropical lowland rainforest is urgently necessary because of the high rate of biodiversity loss and global extinction of species as a result of deforestation. We studied land molluscs species richness and diversity in four heavily degraded secondary forest reserves and one old‐growth forest reserve in Edo State, Nigeria using a combination of direct fixed‐time search and litter‐sieving techniques in twelve plots of 400 m² each per reserve. A total of 43 species and 2570 individuals were collected from all the reserves. Local species richness ranged from 19 to 39 species while number of specimens from 203 to 971. Molluscan species richness and diversity is significantly higher in the old‐growth forest than in the disturbed forest reserves. Land molluscs family composition was similar in all the forests with respect to the common and wide‐ranging species while rare and narrow‐range species are restricted to the old‐growth forest in Okomu. The carnivorous streptaxids and detritivorous subulinids dominate species richness and abundance respectively in all the sites. Species turnover is moderately high within and between the forest reserves indicating the uniqueness of the faunal composition of each forest and the need for adequate protection.     

Allometric constraints and competition enable the simulation of size structure and carbon fluxes in a dynamic vegetation model of tropical forests (LM3PPA‐TV)

Tropical forests are a key determinant of the functioning of the Earth system, but remain a major source of uncertainty in carbon cycle models and climate change projections. In this study, we present an updated land model (LM3PPA‐TV) to improve the representation of tropical forest structure and dynamics in Earth system models (ESMs). The development and parameterization of LM3PPA‐TV drew on extensive datasets on tropical tree traits and long‐term field censuses from Barro Colorado Island (BCI), Panama. The model defines a new plant functional type (PFT) based on the characteristics of shade‐tolerant, tropical tree species, implements a new growth allocation scheme based on realistic tree allometries, incorporates hydraulic constraints on biomass accumulation, and features a new compartment for tree branches and branch fall dynamics. Simulation experiments reproduced observed diurnal and seasonal patterns in stand‐level carbon and water fluxes, as well as mean canopy and understory tree growth rates, tree size distributions, and stand‐level biomass on BCI. Simulations at multiple sites captured considerable variation in biomass and size structure across the tropical forest biome, including observed responses to precipitation and temperature. Model experiments suggested a major role of water limitation in controlling geographic variation forest biomass and structure. However, the failure to simulate tropical forests under extreme conditions and the systematic underestimation of forest biomass in Paleotropical locations highlighted the need to incorporate variation in hydraulic traits and multiple PFTs that capture the distinct floristic composition across tropical domains. The continued pressure on tropical forests from global change demands models which are able to simulate alternative successional pathways and their pace to recovery. LM3PPA‐TV provides a tool to investigate geographic variation in tropical forests and a benchmark to continue improving the representation of tropical forests dynamics and their carbon storage potential in ESMs.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Community divergence in a tropical forest following a severe cyclone

Cyclones are relatively infrequent, may cause massive and widespread disturbance to tropical regions, and are recognized as important determinants of the structure of tropical rainforest communities. Climate change scientists predict that the intensity of cyclones will increase in the future; understanding the long‐term implications of these major disturbances for tropical forest composition and structure will be vital in anticipating and adapting to future changes and impacts. We established a long‐term monitoring site in a rainforest area impacted by severe tropical Cyclone Larry which crossed the North Queensland coast of Australia in March 2006. We monitored recruitment, growth and mortality of nearly 17 000 seedlings in 90 quadrats across the study area for almost 5 years following the cyclone and measured the impact of variation in cyclone disturbance and debris load on community diversity, composition and dispersion as the forest recovered. We show that the level of structural disturbance sustained by the forest has a strong and immediate influence on community dynamics. Quadrats in severely disturbed areas, which were characterized by multiple treefalls and extensive canopy loss, had higher levels of diversity and variation in community assemblage than quadrats in areas characterized primarily by branch loss and defoliation. A rapid divergence in community composition between quadrats in the most‐ and least‐severely disturbed areas resulted in the development of statistically distinct community states across relatively small scales. This provides further evidence that severe cyclones are important in maintaining species diversity in tropical forests.     

Resilience of tropical dry forests – a meta‐analysis of changes in species diversity and composition during secondary succession

Assessing the recovery of species diversity and composition after major disturbance is key to understanding the resilience of tropical forests through successional processes, and its importance for biodiversity conservation. Despite the specific abiotic environment and ecological processes of tropical dry forests, secondary succession has received less attention in this biome than others and changes in species diversity and composition have never been synthesised in a systematic and quantitative review. This study aims to assess in tropical dry forests 1) the directionality of change in species richness and evenness during secondary succession, 2) the convergence of species composition towards that of old‐growth forest and 3) the importance of the previous land use, precipitation regime and water availability in influencing the direction and rate of change. We conducted meta‐analyses of the rate of change in species richness, evenness and composition indices with succession in 13 tropical dry forest chronosequences. Species richness increased with succession, showing a gradual accumulation of species, as did Shannon evenness index. The similarity in species composition of successional forests with old‐growth forests increased with succession, yet at a low rate. Tropical dry forests therefore do show resilience of species composition but it may never reach that of old‐growth forests. We found no significant differences in rates of change between different previous land uses, precipitation regimes or water availability. Our results show high resilience of tropical dry forests in term of species richness but a slow recovery of species composition. They highlight the need for further research on secondary succession in this biome and better understanding of impacts of previous land‐use and landscape‐scale patterns. Synthesis Secondary forests account for an increasing proportion of remaining tropical forest. Assessing their resilience is key to conservation of their biodiversity. Our study is the first meta‐analysis of species changes during succession focussing on tropical dry forests, a highly threatened yet understudied biome. We show a gradual species accumulation and convergence of composition towards that of old‐growth forests. While secondary tropical dry forests offer good potential for biodiversity conservation, their capacity for recovery at a sufficient rate to match threats is uncertain. Further research on this biome is needed to understand the effect of land use history and landscape processes.     

A large‐scale field assessment of carbon stocks in human‐modified tropical forests

Tropical rainforests store enormous amounts of carbon, the protection of which represents a vital component of efforts to mitigate global climate change. Currently, tropical forest conservation, science, policies, and climate mitigation actions focus predominantly on reducing carbon emissions from deforestation alone. However, every year vast areas of the humid tropics are disturbed by selective logging, understory fires, and habitat fragmentation. There is an urgent need to understand the effect of such disturbances on carbon stocks, and how stocks in disturbed forests compare to those found in undisturbed primary forests as well as in regenerating secondary forests. Here, we present the results of the largest field study to date on the impacts of human disturbances on above and belowground carbon stocks in tropical forests. Live vegetation, the largest carbon pool, was extremely sensitive to disturbance: forests that experienced both selective logging and understory fires stored, on average, 40% less aboveground carbon than undisturbed forests and were structurally similar to secondary forests. Edge effects also played an important role in explaining variability in aboveground carbon stocks of disturbed forests. Results indicate a potential rapid recovery of the dead wood and litter carbon pools, while soil stocks (0–30 cm) appeared to be resistant to the effects of logging and fire. Carbon loss and subsequent emissions due to human disturbances remain largely unaccounted for in greenhouse gas inventories, but by comparing our estimates of depleted carbon stocks in disturbed forests with Brazilian government assessments of the total forest area annually disturbed in the Amazon, we show that these emissions could represent up to 40% of the carbon loss from deforestation in the region. We conclude that conservation programs aiming to ensure the long‐term permanence of forest carbon stocks, such as REDD+, will remain limited in their success unless they effectively avoid degradation as well as deforestation.     

Lowland tapirs facilitate seed dispersal in degraded Amazonian forests

The forests of southeastern Amazonia are highly threatened by disturbances such as fragmentation, understory fires, and extreme climatic events. Large‐bodied frugivores such as the lowland tapir (Tapirus terrestris) have the potential to offset this process, supporting natural forest regeneration by dispersing a variety of seeds over long distances to disturbed forests. However, we know little about their effectiveness as seed dispersers in degraded forest landscapes. Here, we investigate the seed dispersal function of lowland tapirs in Amazonian forests subject to a range of human (fire and fragmentation) and natural (extreme droughts and windstorms) disturbances, using a combination of field observations, camera traps, and light detection and ranging (LiDAR) data. Tapirs travel and defecate more often in degraded forests, dispersing much more seeds in these areas [9,822 seeds per ha/year (CI_95% = 9,106; 11,838)] than in undisturbed forests [2,950 seeds per ha/year (CI_95% = 2,961; 3,771)]. By effectively dispersing seeds across disturbed forests, tapirs may contribute to natural forest regeneration—the cheapest and usually the most feasible way to achieve large‐scale restoration of tropical forests. Through the dispersal of large‐seeded species that eventually become large trees, such frugivores also contribute indirectly to maintaining forest carbon stocks. These functions may be critical in helping tropical countries to achieve their goals to maintain and restore biodiversity and its ecosystem services. Ultimately, preserving these animals along with their habitats may help in the process of natural recovery of degraded forests throughout the tropics. Abstract in Portuguese is available with online material.     

Response of an old‐growth tropical rainforest to transient high temperature and drought

Tropical rainforests have experienced episodes of severe heat and drought in recent decades, and climate models project a warmer and potentially drier tropical climate over this century. However, likely responses of tropical rainforests are poorly understood due to a lack of frequent long‐term measurements of forest structure and dynamics. We analyzed a 12‐year record (1999–2010) of 47 817 annual measurements of canopy height to characterize the response of an old‐growth Neotropical rainforest to the severe heat and drought associated with the 1997–1998 El Niño. Well‐drained soils on slopes and plateaus experienced a threefold increase in the fraction of the landscape in gaps (≤2 m) and a reduction in the fraction in high canopy (>15 m) causing distributions of canopy height to depart from equilibrium for a period of 2–3 years. In contrast, forests on low‐lying alluvial terraces remained in equilibrium and were nearly half as likely to experience upper canopy (>15 m) disturbance over the 12 years of observation. Variation in forest response across topographic positions suggests that tropical rainforests are more sensitive to moisture deficits than high temperature and that topography likely structures landscape‐level variation in the severity of drought impacts.     

Gap formation and dynamics after long‐term steady state in an old‐growth Picea abies stand in Norway: Above‐ and belowground interactions

Stand dynamics and the gap initiation prior to gap formation are not well‐understood because of its long‐term nature and the scarcity of late‐successional stands. Reconstruction of such disturbance is normally based on historical records and dendroecological methods. We investigated gap initiation and formation at the fine‐scale stand level in the old‐growth reserve of Karlshaugen in Norway. Given its long‐term conservation history, and thorough mapping in permanent marked plots with spatially referenced trees, it provides an opportunity to present stand development before, during, and after gap formation. Late‐successional decline in biomass was recorded after more than 50 years of close to steady state. Gaps in the canopy were mainly created by large old trees that had been killed by spruce bark beetles. Snapping by wind was the main reason for treefall. Long‐term dominance of Norway spruce excluded downy birch and Scots pine from the stand. Comparisons of the forest floor soil properties between the gap and nongap area showed significantly higher concentrations of plant available Ca within the gap area. Plant root simulator (PRS?) probes showed significantly higher supply rates for Ca and Mg, but significantly lower K for the gap compared to the nongap area. Soil water from the gap area had significantly higher C:N ratios compared to the nongap area. Fine‐scale variation with increasing distance to logs indicated that CWD is important for leaking of DOC and Ca. Our long‐term study from Karlshaugen documents gap dynamics after more than 50 years of steady state and a multiscale disturbance regime in an old‐growth forest. The observed disturbance dynamic caused higher aboveground and belowground heterogeneity in plots, coarse woody debris, and nutrients. Our study of the nutrient levels of the forest floor suggest that natural gaps of old‐growth forest provide a long‐lasting biogeochemical feedback system particularly with respect to Ca and probably also N. Norway spruce trees near the gap edge responded with high plasticity to reduced competition, showing the importance of the edge zone as hot spots for establishing heterogeneity, but also the potential for carbon sequestration in old‐growth forest.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

Coordinated approaches to quantify long‐term ecosystem dynamics in response to global change

Many serious ecosystem consequences of climate change will take decades or even centuries to emerge. Long‐term ecological responses to global change are strongly regulated by slow processes, such as changes in species composition, carbon dynamics in soil and by long‐lived plants, and accumulation of nutrient capitals. Understanding and predicting these processes require experiments on decadal time scales. But decadal experiments by themselves may not be adequate because many of the slow processes have characteristic time scales much longer than experiments can be maintained. This article promotes a coordinated approach that combines long‐term, large‐scale global change experiments with process studies and modeling. Long‐term global change manipulative experiments, especially in high‐priority ecosystems such as tropical forests and high‐latitude regions, are essential to maximize information gain concerning future states of the earth system. The long‐term experiments should be conducted in tandem with complementary process studies, such as those using model ecosystems, species replacements, laboratory incubations, isotope tracers, and greenhouse facilities. Models are essential to assimilate data from long‐term experiments and process studies together with information from long‐term observations, surveys, and space‐for‐time studies along environmental and biological gradients. Future research programs with coordinated long‐term experiments, process studies, and modeling have the potential to be the most effective strategy to gain the best information on long‐term ecosystem dynamics in response to global change.     

Ecological outcomes and livelihood benefits of community‐managed agroforests and second growth forests in Southeast Brazil

The Forest and Landscape Restoration movement has emerged as an approach to reconcile biodiversity conservation, ecosystem services provisioning and human well‐being in degraded landscapes, but little is known so far about the potential of different reforestation methods to achieve these objectives. Based on this gap, we assessed the ecological outcomes and local livelihood benefits of community‐managed agroforests and second growth forests to assist natural regeneration in the coastal Atlantic Forest of Brazil. We investigated and compared agroforests and secondary forests according to their structure and floristic composition in 51 circular plots of 314 m², their role in supporting local livelihoods (45 semi‐structured interviews) and the use and cultural importance of plant species (61 interviews). Agroforests and, more remarkably, managed secondary forests (1) re‐established a well‐developed forest structure, with a higher density of tree‐sized individuals and similar basal area compared to nearby old growth forests; (2) were composed by a rich array of native species, including five threatened species, but had lower species richness than old growth remnants; and (3) improved local livelihoods by supplying market valuable and culturally important plants, including 231 native ethnospecies. Overall, local production systems showed remarkable potential to engage smallholders of developing tropical countries in Forest and Landscape Restoration and contribute to achieve its overall goals. We advocate the promotion of these systems as effective Forest and Landscape Restoration approaches in multi‐scale programs and policies.     

Forest Regeneration in a Chronosequence of Tropical Abandoned Pastures: Implications for Restoration Ecology

During the mid‐1900s, most of the island of Puerto Rico was deforested, but a shift in the economy from agriculture to small industry beginning in the 1950s resulted in the abandonment of agricultural lands and recovery of secondary forest. This unique history provides an excellent opportunity to study secondary forest succession and suggest strategies for tropical forest restoration. To determine the pattern of secondary succession, we describe the woody vegetation in 71 abandoned pastures and forest sites in four regions of Puerto Rico. The density, basal area, aboveground biomass, and species richness of the secondary forest sites were similar to those of the old growth forest sites (>80 yr) after approximately 40 years. The dominant species that colonized recently abandoned pastures occurred over a broad elevational range and are widespread in the neotropics. The species richness of Puerto Rican secondary forests recovered rapidly, but the species composition was quite different in comparison with old growth forest sites, suggesting that enrichment planting will be necessary to restore the original composition. Exotic species were some of the most abundant species in the secondary forest, but their long‐term impact depended on life history characteristics of each species. These data demonstrate that one restoration strategy for tropical forest in abandoned pastures is simply to protect the areas from fire, and allow natural regeneration to produce secondary forest. This strategy will be most effective if remnant forest (i.e., seed sources) still exist in the landscape and soils have not been highly degraded. Patterns of forest recovery also suggest strategies for accelerating natural recovery by planting a suite of generalist species that are common in recently abandoned pastures in Puerto Rico and throughout much of the neotropics.     

Long‐term recovery of the functional community assembly and carbon pools in an African tropical forest succession

On the African continent, the population is expected to expand fourfold in the next century, which will increasingly impact the global carbon cycle and biodiversity conservation. Therefore, it is of vital importance to understand how carbon stocks and community assembly recover after slash‐and‐burn events in tropical second growth forests. We inventoried a chronosequence of 15 1‐ha plots in lowland tropical forest of the central Congo Basin and evaluated changes in aboveground and soil organic carbon stocks and in tree species diversity, functional composition, and community‐weighted functional traits with succession. We aimed to track long‐term recovery trajectories of species and carbon stocks in secondary forests, comparing 5 to 200 + year old secondary forest with reference primary forest. Along the successional gradient, the functional composition followed a trajectory from resource acquisition to resource conservation, except for nitrogen‐related leaf traits. Despite a fast, initial recovery of species diversity and functional composition, there were still important structural and carbon stock differences between old growth secondary and pristine forest, which suggests that a full recovery of secondary forests might take much longer than currently shown. As such, the aboveground carbon stocks of 200 + year old forest were only 57% of those in the pristine reference forest, which suggests a slow recovery of aboveground carbon stocks, although more research is needed to confirm this observation. The results of this study highlight the need for more in‐depth studies on forest recovery in Central Africa, to gain insight into the processes that control biodiversity and carbon stock recovery.     

Multiple successional pathways in human‐modified tropical landscapes: new insights from forest succession,forest fragmentation and landscape ecology research

Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.