Light competition drives species replacement during secondary tropical forest succession.

Light competition is thought to drive successional shifts in species dominance in closed vegetations, but few studies have assessed this for species-rich and vertically structured tropical forests. We analyzed how light competition drives species replacement during succession, and how cross-species variation in light competition strategies is determined by underlying species traits. To do so, we used chronosequence approach in which we compared 14 Mexican tropical secondary rainforest stands that differ in age (8-32 year-old). For each tree, height and stem diameter were monitored for 2 years to calculate relative biomass growth rate (RGR, the aboveground biomass gain per unit aboveground tree biomass per year). For each stand, 3D light profiles were measured to estimate individuals' light interception to calculate light interception efficiency (LIE, intercepted light per unit biomass per year) and light use efficiency (LUE, biomass growth per intercepted light). Throughout succession, species with higher RGR attained higher changes in species dominance and thus increased their dominance over time. Both light competition strategies (LIE and LUE) increased RGR. In early succession, a high LIE and its associated traits (large crown leaf mass and low wood density) are more important for RGR. During succession, forest structure builds up, leading to lower understory light levels. In later succession, a high LUE and its associated traits (high wood density and leaf mass per area) become more important for RGR. Therefore, successional changes in relative importance of light competition strategies drive shifts in species dominance during tropical rainforest succession.

Functional recovery of secondary tropical forests.

One-third of all Neotropical forests are secondary forests that regrow naturally after agricultural use through secondary succession. We need to understand better how and why succession varies across environmental gradients and broad geographic scales. Here, we analyze functional recovery using community data on seven plant characteristics (traits) of 1,016 forest plots from 30 chronosequence sites across the Neotropics. By analyzing communities in terms of their traits, we enhance understanding of the mechanisms of succession, assess ecosystem recovery, and use these insights to propose successful forest restoration strategies. Wet and dry forests diverged markedly for several traits that increase growth rate in wet forests but come at the expense of reduced drought tolerance, delay, or avoidance, which is important in seasonally dry forests. Dry and wet forests showed different successional pathways for several traits. In dry forests, species turnover is driven by drought tolerance traits that are important early in succession and in wet forests by shade tolerance traits that are important later in succession. In both forests, deciduous and compound-leaved trees decreased with forest age, probably because microclimatic conditions became less hot and dry. Our results suggest that climatic water availability drives functional recovery by influencing the start and trajectory of succession, resulting in a convergence of community trait values with forest age when vegetation cover builds up. Within plots, the range in functional trait values increased with age. Based on the observed successional trait changes, we indicate the consequences for carbon and nutrient cycling and propose an ecologically sound strategy to improve forest restoration success.

Landscape-scale forest cover drives the predictability of forest regeneration across the Neotropics.

Abandonment of agricultural lands promotes the global expansion of secondary forests, which are critical for preserving biodiversity and ecosystem functions and services. Such roles largely depend, however, on two essential successional attributes, trajectory and recovery rate, which are expected to depend on landscape-scale forest cover in nonlinear ways. Using a multi-scale approach and a large vegetation dataset (843 plots, 3511 tree species) from 22 secondary forest chronosequences distributed across the Neotropics, we show that successional trajectories of woody plant species richness, stem density and basal area are less predictable in landscapes (4 km radius) with intermediate (40-60%) forest cover than in landscapes with high (greater than 60%) forest cover. This supports theory suggesting that high spatial and environmental heterogeneity in intermediately deforested landscapes can increase the variation of key ecological factors for forest recovery (e.g. seed dispersal and seedling recruitment), increasing the uncertainty of successional trajectories. Regarding the recovery rate, only species richness is positively related to forest cover in relatively small (1 km radius) landscapes. These findings highlight the importance of using a spatially explicit landscape approach in restoration initiatives and suggest that these initiatives can be more effective in more forested landscapes, especially if implemented across spatial extents of 1-4 km radius.

Biomass resilience of Neotropical secondary forests.

Land-use change occurs nowhere more rapidly than in the tropics, where the imbalance between deforestation and forest regrowth has large consequences for the global carbon cycle. However, considerable uncertainty remains about the rate of biomass recovery in secondary forests, and how these rates are influenced by climate, landscape, and prior land use. Here we analyse aboveground biomass recovery during secondary succession in 45 forest sites and about 1,500 forest plots covering the major environmental gradients in the Neotropics. The studied secondary forests are highly productive and resilient. Aboveground biomass recovery after 20 years was on average 122 megagrams per hectare (Mg ha(-1)), corresponding to a net carbon uptake of 3.05 Mg C ha(-1) yr(-1), 11 times the uptake rate of old-growth forests. Aboveground biomass stocks took a median time of 66 years to recover to 90% of old-growth values. Aboveground biomass recovery after 20 years varied 11.3-fold (from 20 to 225 Mg ha(-1)) across sites, and this recovery increased with water availability (higher local rainfall and lower climatic water deficit). We present a biomass recovery map of Latin America, which illustrates geographical and climatic variation in carbon sequestration potential during forest regrowth. The map will support policies to minimize forest loss in areas where biomass resilience is naturally low (such as seasonally dry forest regions) and promote forest regeneration and restoration in humid tropical lowland areas with high biomass resilience.

Anthropogenic disturbances jeopardize biodiversity conservation within tropical rainforest reserves.

Anthropogenic disturbances affecting tropical forest reserves have been documented, but their ecological long-term cumulative effects are poorly understood. Habitat fragmentation and defaunation are two major anthropogenic threats to the integrity of tropical reserves. Based on a long-term (four decades) study, we document how these disturbances synergistically disrupt ecological processes and imperil biodiversity conservation and ecosystem functioning at Los Tuxtlas, the northernmost tropical rainforest reserve in the Americas. Deforestation around this reserve has reduced the reserve to a medium-sized fragment (640 ha), leading to an increased frequency of canopy-gap formation. In addition, hunting and habitat loss have caused the decline or local extinction of medium and large herbivores. Combining empirical, experimental, and modeling approaches, we support the hypothesis that such disturbances produced a demographic explosion of the long-lived (≈120 y old, maximum height of 7 m) understory palm Astrocaryum mexicanum, whose population has increased from 1,243-4,058 adult individuals per hectare in only 39 y (annual growth rate of ca 3%). Faster gap formation increased understory light availability, enhancing seed production and the growth of immature palms, whereas release from mammalian herbivory and trampling increased survival of seedlings and juveniles. In turn, the palm's demographic explosion was followed by a reduction of tree species diversity, changing forest composition, altering the relative contribution of trees to forest biomass, and disrupting litterfall dynamics. We highlight how indirect anthropogenic disturbances (e.g., palm proliferation) on otherwise protected areas threaten tropical conservation, a phenomenon that is currently eroding the planet's richest repositories of biodiversity.

Fragmentation and matrix contrast favor understory plants through negative cascading effects on a strong competitor palm.

Understanding the patterns and processes driving biodiversity maintenance in fragmented tropical forests is urgently needed for conservation planning, especially in species-rich forest reserves. Of particular concern are the effects that habitat modifications at the landscape scale may have on forest regeneration and ecosystem functioning: a topic that has received limited attention. Here, we assessed the effects of landscape structure (i.e., forest cover, open area matrices, forest fragmentation, and mean inter-patch isolation distance) on understory plant assemblages in the Los Tuxtlas Biosphere Reserve, Mexico. Previous studies suggest that the demographic burst of the strong competitor palm Astrocaryum mexicanum in the core area of this reserve limits plant recruitment and imperils biodiversity conservation within this protected area. Yet, the local and landscape predictors of this palm, and its impact on tree recruitment at a regional scale are unknown. Thus, we used structural equation modeling to assess the direct and cascading effects of landscape structure on stem and species density in the understory of 20 forest sites distributed across this biodiversity hotspot. Indirect paths included the effect of landscape structure on tree basal area (a proxy of local disturbance), and the effects of these variables on A. mexicanum. Density of A. mexicanum mainly increased with decreasing both fragmentation and open areas in the matrix (matrix contrast, hereafter), and such an increase in palm density negatively affected stem and species density in the understory. The negative direct effect of matrix contrast on stem density was overridden by the indirect positive effects (i.e., through negative cascading effects on A. mexicanum), resulting in a weak effect of matrix contrast on stem density. These findings suggest that dispersal limitation and negative edge effects in more fragmented landscapes dominated by open areas prevent the proliferation of this palm species, enhancing the diversity and abundance of understory trees. This "positive" news adds to an increasing line of evidence suggesting that fragmentation may have some positive effects on biodiversity, in this case by preventing the proliferation of species that can jeopardize biodiversity conservation within tropical reserves.

Successional dynamics in Neotropical forests are as uncertain as they are predictable.

Although forest succession has traditionally been approached as a deterministic process, successional trajectories of vegetation change vary widely, even among nearby stands with similar environmental conditions and disturbance histories. Here, we provide the first attempt, to our knowledge, to quantify predictability and uncertainty during succession based on the most extensive long-term datasets ever assembled for Neotropical forests. We develop a novel approach that integrates deterministic and stochastic components into different candidate models describing the dynamical interactions among three widely used and interrelated forest attributes--stem density, basal area, and species density. Within each of the seven study sites, successional trajectories were highly idiosyncratic, even when controlling for prior land use, environment, and initial conditions in these attributes. Plot factors were far more important than stand age in explaining successional trajectories. For each site, the best-fit model was able to capture the complete set of time series in certain attributes only when both the deterministic and stochastic components were set to similar magnitudes. Surprisingly, predictability of stem density, basal area, and species density did not show consistent trends across attributes, study sites, or land use history, and was independent of plot size and time series length. The model developed here represents the best approach, to date, for characterizing autogenic successional dynamics and demonstrates the low predictability of successional trajectories. These high levels of uncertainty suggest that the impacts of allogenic factors on rates of change during tropical forest succession are far more pervasive than previously thought, challenging the way ecologists view and investigate forest regeneration.

Demographic drivers of functional composition dynamics.

Mechanisms of community assembly and ecosystem function are often analyzed using community-weighted mean trait values (CWMs). We present a novel conceptual framework to quantify the contribution of demographic processes (i.e., growth, recruitment, and mortality) to temporal changes in CWMs. We used this framework to analyze mechanisms of secondary succession in wet tropical forests in Mexico. Seed size increased over time, reflecting a trade-off between colonization by small seeds early in succession, to establishment by large seeds later in succession. Specific leaf area (SLA) and leaf phosphorus content decreased over time, reflecting a trade-off between fast growth early in succession vs. high survival late in succession. On average, CWM shifts were driven mainly (70%) by growth of surviving trees that comprise the bulk of standing biomass, then mortality (25%), and weakly by recruitment (5%). Trait shifts of growing and recruiting trees mirrored the CWM trait shifts, and traits of dying trees did not change during succession, indicating that these traits are important for recruitment and growth, but not for mortality, during the first 30 yr of succession. Identifying the demographic drivers of functional composition change links population dynamics to community change, and enhances insights into mechanisms of succession.

Explaining long-term inter-individual growth variation in plant populations: persistence of abiotic factors matters.

An unanswered question in ecology is whether the environmental factors driving short-term performance also determine the often observed long-term performance differences among individuals. Here, we analyze the extent to which temporal persistence of spatial heterogeneity in environmental factors can contribute to long-term inter-individual variation in stem length growth. For a natural population of a long-lived understorey palm, we first quantified the effect of several environmental factors on stem length growth and survival. We then performed individual-based simulations of growth trajectories, in which we varied, for two environmental factors: (1) the strength of the effect on stem length growth and (2) the temporal persistence. Short-term variation in stem length growth was strongly driven by light availability. Auto-correlation in light availability and soil pH increased simulated variation in stem length growth among 20-year-old palms to levels similar to the observed variation. Analyses in which we varied both the strength of the effect on stem length growth and the temporal persistence of the environmental factors revealed that a large fraction of observed long-term growth differences can be explained, as long as one of these effects is strong. This implies that environmental factors that are relatively unimportant for short-term performance can still drive long-term performance differences when the environmental variation is sufficiently persistent over time.

The importance of biodiversity and dominance for multiple ecosystem functions in a human-modified tropical landscape.

Many studies suggest that biodiversity may be particularly important for ecosystem multifunctionality, because different species with different traits can contribute to different functions. Support, however, comes mostly from experimental studies conducted at small spatial scales in low-diversity systems. Here, we test whether different species contribute to different ecosystem functions that are important for carbon cycling in a high-diversity human-modified tropical forest landscape in Southern Mexico. We quantified aboveground standing biomass, primary productivity, litter production, and wood decomposition at the landscape level, and evaluated the extent to which tree species contribute to these ecosystem functions. We used simulations to tease apart the effects of species richness, species dominance and species functional traits on ecosystem functions. We found that dominance was more important than species traits in determining a species' contribution to ecosystem functions. As a consequence of the high dominance in human-modified landscapes, the same small subset of species mattered across different functions. In human-modified landscapes in the tropics, biodiversity may play a limited role for ecosystem multifunctionality due to the potentially large effect of species dominance on biogeochemical functions. However, given the spatial and temporal turnover in species dominance, biodiversity may be critically important for the maintenance and resilience of ecosystem functions.

Experimental defaunation of terrestrial mammalian herbivores alters tropical rainforest understorey diversity.

It has been suggested that tropical defaunation may unleash community-wide cascading effects, leading to reductions in plant diversity. However, experimental evidence establishing cause-effect relationships thereof is poor. Through a 5 year exclosure experiment, we tested the hypothesis that mammalian defaunation affects tree seedling/sapling community dynamics leading to reductions in understorey plant diversity. We established plot triplets (n = 25) representing three defaunation contexts: terrestrial-mammal exclosure (TE), medium/large mammal exclosure (PE) and open access controls (C). Seedlings/saplings 30-100 cm tall were marked and identified within each of these plots and re-censused three times to record survival and recruitment. In the periods 2010-2011 and 2011-2013, survival was greater in PE than in C plots and recruitment was higher in TE plots than in C plots. Overall, seedling density increased by 61% in TE plots and 23% in PE plots, whereas it decreased by 5% in C plots. Common species highly consumed by mammals (e.g. Brosimum alicastrum and Ampelocera hottlei) increased in their abundance in TE plots. Rarefaction curves showed that species diversity decreased in TE plots from 2008 to 2013, whereas it remained similar for C plots. Given the prevalence of tropical defaunation, we posit this is an anthropogenic effect threatening the maintenance of tropical forest diversity.

Biomass is the main driver of changes in ecosystem process rates during tropical forest succession.

Over half of the world's forests are disturbed, and the rate at which ecosystem processes recover after disturbance is important for the services these forests can provide. We analyze the drivers' underlying changes in rates of key ecosystem processes (biomass productivity, litter productivity, actual litter decomposition, and potential litter decomposition) during secondary succession after shifting cultivation in wet tropical forest of Mexico. We test the importance of three alternative drivers of ecosystem processes: vegetation biomass (vegetation quantity hypothesis), community-weighted trait mean (mass ratio hypothesis), and functional diversity (niche complementarity hypothesis) using structural equation modeling. This allows us to infer the relative importance of different mechanisms underlying ecosystem process recovery. Ecosystem process rates changed during succession, and the strongest driver was aboveground biomass for each of the processes. Productivity of aboveground stem biomass and leaf litter as well as actual litter decomposition increased with initial standing vegetation biomass, whereas potential litter decomposition decreased with standing biomass. Additionally, biomass productivity was positively affected by community-weighted mean of specific leaf area, and potential decomposition was positively affected by functional divergence, and negatively by community-weighted mean of leaf dry matter content. Our empirical results show that functional diversity and community-weighted means are of secondary importance for explaining changes in ecosystem process rates during tropical forest succession. Instead, simply, the amount of vegetation in a site is the major driver of changes, perhaps because there is a steep biomass buildup during succession that overrides more subtle effects of community functional properties on ecosystem processes. We recommend future studies in the field of biodiversity and ecosystem functioning to separate the effects of vegetation quality (community-weighted mean trait values and functional diversity) from those of vegetation quantity (biomass) on ecosystem processes and services.

Primate extirpation from rainforest fragments does not appear to influence seedling recruitment.

Primates are important seed dispersers, especially of large-seeded tree species, but the impact that these animals have on seedling recruitment is unclear. Evidence suggests that forest regeneration might be disrupted in forest fragments in which primates were extirpated. We tested this hypothesis by assessing seedling recruitment in 3 forest fragments occupied (OF) by primates, 3 fragments unoccupied (UF) by primates, and 3 areas within a continuous forest (CF) in the Lacandona rainforest, Mexico. Species and stem densities of tree, palm and liana seedlings were recorded over 16 months. Individuals were classified according to dispersal mode: large-seeded animal-dispersed (LS), small- and medium-seeded animal-dispersed (SS), and abiotically-dispersed species (AD). We assessed the influence of primate presence, adult tree assemblage, and fragment spatial metrics (size, age, distance to nearest fragment, and distance to continuous forest) on seedling assemblages. We recorded 6879 seedlings belonging to 90 species, and 59 genera in 405 1-m(2) plots. Both seedlings and adults showed similar differences in species and stem densities of LS, SS and AD species among forest types, suggesting that seedling assemblages were strongly influenced by the adult assemblages. The recruitment of each LS species varied among study sites, but evidence supporting higher recruitment enhancement of these species in continuous forest and occupied fragments was weak. Distance to continuous forest was the unique fragment spatial metric related (negatively) to the recruitment of LS species. Thus, primate extirpation does not appear to disrupt seedling assemblages in the Lancandona rainforest. Nevertheless, we cannot reject the hypothesis that certain LS species such as Spondias radlkoferi may be affected by the extirpation of primates.

Feedback loops drive ecological succession: towards a unified conceptual framework.

The core principle shared by most theories and models of succession is that, following a major disturbance, plant-environment feedback dynamics drive a directional change in the plant community. The most commonly studied feedback loops are those in which the regrowth of the plant community causes changes to the abiotic (e.g. soil nutrients) or biotic (e.g. dispersers) environment, which differentially affect species availability or performance. This, in turn, leads to shifts in the species composition of the plant community. However, there are many other PE feedback loops that potentially drive succession, each of which can be considered a model of succession. While plant-environment feedback loops in principle generate predictable successional trajectories, succession is generally observed to be highly variable. Factors contributing to this variability are the stochastic processes involved in feedback dynamics, such as individual mortality and seed dispersal, and extrinsic causes of succession, which are not affected by changes in the plant community but do affect species performance or availability. Both can lead to variation in the identity of dominant species within communities. This, in turn, leads to further contingencies if these species differ in their effect on their environment (priority effects). Predictability and variability are thus intrinsically linked features of ecological succession. We present a new conceptual framework of ecological succession that integrates the propositions discussed above. This framework defines seven general causes: landscape context, disturbance and land-use, biotic factors, abiotic factors, species availability, species performance, and the plant community. When involved in a feedback loop, these general causes drive succession and when not, they are extrinsic causes that create variability in successional trajectories and dynamics. The proposed framework provides a guide for linking these general causes into causal pathways that represent specific models of succession. Our framework represents a systematic approach to identifying the main feedback processes and causes of variation at different successional stages. It can be used for systematic comparisons among study sites and along environmental gradients, to conceptualise studies, and to guide the formulation of research questions and design of field studies. Mapping an extensive field study onto our conceptual framework revealed that the pathways representing the study's empirical outcomes and conceptual model had important differences, underlining the need to move beyond the conceptual models that currently dominate in specific fields and to find ways to examine the importance of and interactions among alternative causal pathways of succession. To further this aim, we argue for integrating long-term studies across environmental and anthropogenic gradients, combined with controlled experiments and dynamic modelling.

Correlations between physical and chemical defences in plants: tradeoffs, syndromes, or just many different ways to skin a herbivorous cat?

Most plant species have a range of traits that deter herbivores. However, understanding of how different defences are related to one another is surprisingly weak. Many authors argue that defence traits trade off against one another, while others argue that they form coordinated defence syndromes. We collected a dataset of unprecedented taxonomic and geographic scope (261 species spanning 80 families, from 75 sites across the globe) to investigate relationships among four chemical and six physical defences. Five of the 45 pairwise correlations between defence traits were significant and three of these were tradeoffs. The relationship between species' overall chemical and physical defence levels was marginally nonsignificant (P = 0.08), and remained nonsignificant after accounting for phylogeny, growth form and abundance. Neither categorical principal component analysis (PCA) nor hierarchical cluster analysis supported the idea that species displayed defence syndromes. Our results do not support arguments for tradeoffs or for coordinated defence syndromes. Rather, plants display a range of combinations of defence traits. We suggest this lack of consistent defence syndromes may be adaptive, resulting from selective pressure to deploy a different combination of defences to coexisting species.

Successional changes in functional composition contrast for dry and wet tropical forest.

We tested whether and how functional composition changes with succession in dry deciduous and wet evergreen forests of Mexico. We hypothesized that compositional changes during succession in dry forest were mainly determined by increasing water availability leading to community functional changes from conservative to acquisitive strategies, and in wet forest by decreasing light availability leading to changes from acquisitive to conservative strategies. Research was carried out in 15 dry secondary forest plots (5-63 years after abandonment) and 17 wet secondary forest plots (< 1-25 years after abandonment). Community-level functional traits were represented by community-weighted means based on 11 functional traits measured on 132 species. Successional changes in functional composition are more marked in dry forest than in wet forest and largely characterized by different traits. During dry forest succession, conservative traits related to drought tolerance and drought avoidance decreased, as predicted. Unexpectedly acquisitive leaf traits also decreased, whereas seed size and dependence on biotic dispersal increased. In wet forest succession, functional composition changed from acquisitive to conservative leaf traits, suggesting light availability as the main driver of changes. Distinct suites of traits shape functional composition changes in dry and wet forest succession, responding to different environmental filters.

Successional theories.

Succession is a fundamental concept in ecology because it indicates how species populations, communities, and ecosystems change over time on new substrate or after a disturbance. A mechanistic understanding of succession is needed to predict how ecosystems will respond to land-use change and to design effective ecosystem restoration strategies. Yet, despite a century of conceptual advances a comprehensive successional theory is lacking. Here we provide an overview of 19 successional theories ('models') and their key points, group them based on conceptual similarity, explain conceptual development in successional ideas and provide suggestions how to move forward. Four groups of models can be recognised. The first group (patch & plants) focuses on plants at the patch level and consists of three subgroups that originated in the early 20th century. One subgroup focuses on the processes (dispersal, establishment, and performance) that operate sequentially during succession. Another subgroup emphasises individualistic species responses during succession, and how this is driven by species traits. A last subgroup focuses on how vegetation structure and underlying demographic processes change during succession. A second group of models (ecosystems) provides a more holistic view of succession by considering the ecosystem, its biota, interactions, diversity, and ecosystem structure and processes. The third group (landscape) considers a larger spatial scale and includes the effect of the surrounding landscape matrix on succession as the distance to neighbouring vegetation patches determines the potential for seed dispersal, and the quality of the neighbouring patches determines the abundance and composition of seed sources and biotic dispersal vectors. A fourth group (socio-ecological systems) includes the human component by focusing on socio-ecological systems where management practices have long-lasting legacies on successional pathways and where regrowing vegetations deliver a range of ecosystem services to local and global stakeholders. The four groups of models differ in spatial scale (patch, landscape) or organisational level (plant species, ecosystem, socio-ecological system), increase in scale and scope, and reflect the increasingly broader perspective on succession over time. They coincide approximately with four periods that reflect the prevailing view of succession of that time, although all views still coexist. The four successional views are: succession of plants (from 1910 onwards) where succession was seen through the lens of species replacement; succession of communities and ecosystems (from 1965 onwards) when there was a more holistic view of succession; succession in landscapes (from 2000 onwards) when it was realised that the structure and composition of landscapes strongly impact successional pathways, and increased remote-sensing technology allowed for a better quantification of the landscape context; and succession with people (from 2015 onwards) when it was realised that people and societal drivers have strong effects on successional pathways, that ecosystem processes and services are important for human well-being, and that restoration is most successful when it is done by and for local people. Our review suggests that the hierarchical successional framework of Pickett is the best starting point to move forward as this framework already includes several factors, and because it is flexible, enabling application to different systems. The framework focuses mainly on species replacement and could be improved by focusing on succession occurring at different hierarchical scales (population, community, ecosystem, socio-ecological system), and by integrating it with more recent developments and other successional models: by considering different spatial scales (landscape, region), temporal scales (ecosystem processes occurring over centuries, and evolution), and by taking the effects of the surrounding landscape (landscape integrity and composition, the disperser community) and societal factors (previous and current land-use intensity) into account. Such a new, comprehensive framework could be tested using a combination of empirical research, experiments, process-based modelling and novel tools. Applying the framework to seres across broadscale environmental and disturbance gradients allows a better insight into what successional processes matter and under what conditions.

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes.

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Forest structure drives changes in light heterogeneity during tropical secondary forest succession.

Light is a key resource for tree performance and hence, tree species partition spatial and temporal gradients in light availability. Although light distribution drives tree performance and species replacement during secondary forest succession, we yet lack understanding how light distribution changes with tropical forest development.This study aims to evaluate how changes in forest structure lead to changes in vertical and horizontal light heterogeneity during tropical forest succession.We described successional patterns in light using a chronosequence approach in which we compared 14 Mexican secondary forest stands that differ in age (8-32 years) since agricultural abandonment. For each stand, we measured vertical light profiles in 16 grid cells, and structural parameters (diameter at breast height, height and crown dimensions) for each tree.During succession, we found a rapid increase in stand size (basal area, crown area and length) and stand differentiation (i.e. a gradual leaf distribution along the forest profile), which leads to fast changes in light conditions and more light heterogeneity. The inflection points of the vertical light gradient (i.e. the absolute height at which 50% relative light intensity is attained) rapidly moved towards higher heights in the first 20 years, indicating that larger amounts of light are intercepted by canopy trees. Light attenuation rate (i.e. the rate of light extinction) decreased during succession due to slower accumulation of the crown area with height. Understorey light intensity and heterogeneity slightly decreased during succession because of an increase in crown size and a decrease in lateral gap frequency. Understorey relative light intensity was 1.56% at 32 years after abandonment.Synthesis. During succession, light conditions changed linearly, which should lead to a continuous and constant replacement of species. Especially in later successional stages, stronger vertical light gradients can limit the regeneration of light-demanding pioneer species and increase the proportion of shade-tolerant late-successional species under the canopy. These changes in light conditions were largely driven by the successional changes in forest structure, as basal area strongly determined the height where most light is absorbed, whereas crown area, and to a lesser extent crown length, determined light distribution.

Multidimensional tropical forest recovery.

Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and slowest for biomass and species composition (>12 decades). Network analysis shows three independent clusters of attribute recovery, related to structure, species diversity, and species composition. Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration, climate change mitigation, and biodiversity conservation.