Tropical Forest Succession Research Articles

Soil fungi lead to stronger ‘diminishing returns’ in fine‐root length versus mass allometry towards earlier successional tropical forests

Soil fungi lead to stronger ‘diminishing returns’ in fine‐root length versus mass allometry towards earlier successional tropical forests

Combining spaceborne lidar from the Global Ecosystem Dynamics Investigation with local knowledge for monitoring fragmented tropical landscapes: A case study in the forest-agriculture interface of Ucayali, Peru.

Improving our ability to monitor fragmented tropical ecosystems is a critical step in supporting the stewardship of these complex landscapes. We investigated the structural characteristics of vegetation classes in Ucayali, Peru, employing a co-production approach. The vegetation classes included three agricultural classes (mature oil palm, monocrop cacao, and agroforestry cacao plantations) and three forest regeneration classes (mature lowland forest, secondary lowland forest, and young lowland vegetation regrowth). We combined local knowledge with spaceborne lidar from NASA's Global Ecosystem Dynamics Investigation mission to classify vegetation and characterize the horizontal and vertical structure of each vegetation class. Mature lowland forest had consistently higher mean canopy height and lower canopy height variance than secondary lowland forest (μ = 29.40 m, sd = 6.89 m vs. μ = 20.82 m, sd = 9.15 m, respectively). The lower variance of mature forest could be attributed to the range of forest development ages in the secondary forest patches. However, secondary forests exhibited a similar vertical profile to mature forests, with each cumulative energy percentile increasing at similar rates. We also observed similar mean and standard deviations in relative height ratios (RH50/RH95) for mature forest, secondary forest, and oil palm even when removing the negative values from the relative height ratios and interpolating from above-ground returns only (mean RH50/RH95 of 0.58, 0.54, and 0.53 for mature forest, secondary forest, and oil palm, respectively) (p < .0001). This pattern differed from our original expectations based on local knowledge and existing tropical forest succession studies, pointing to opportunities for future work. Our findings suggest that lidar-based relative height metrics can complement local information and other remote sensing approaches that rely on optical imagery, which are limited by extensive cloud cover in the tropics. We show that characterizing ecosystem structure with a co-production approach can support addressing both the technical and social challenges of monitoring and managing fragmented tropical landscapes.

Diversity and Distribution of Monocot Understory Herbs during Tropical Forest Succession in Northeastern Costa Rica

Broad-leaved monocot herbs form one of the most common and diverse growth forms of Neotropical plants. Their significance and frequency of occurrence is particularly notable in the understories of tropical rainforests, where they form a dominant element. We assessed and quantified changes in the cover and diversity of understory herb communities in a chronosequence of 1 ha permanent plots established as part of a multidisciplinary study on tropical forest regeneration in the Atlantic lowlands of northeastern Costa Rica. Sampled were two young stands cleared 12 years ago, two secondary forests with 21 and 39 of years of recovery since clearance, and two stands in old-growth primary forest. Changes in species composition during succession were assessed using Chao’s Jaccard similarity index. Observed species richness ranged from 15 to 26 species in individual plots, with the greatest number of species in the 21-year intermediate-age and fewest in the young 12-year plots. Herb species sampled represented 6 families, 15 genera, and 39 species, with the Araceae contributing the largest number of species. Ten species were sampled in all six stands, while fourteen species were found exclusively in one plot. Herb density (ramets m−2) showed a hump-shade trend, with peak density in the intermediate stands and a lower level in mature and young secondary forests. Mean herb cover in 25 m2 quadrats ranged from 2.0% (young stand) to 22.7% (intermediate-age stand) and differed significantly both among stand types and among sites. Both observed and estimated species richness increased along the chronosequence as a whole, with the highest number of species in primary forest, although only slightly higher than in intermediate-age stands. Over half of the species exhibited some degree of clonal growth, with the extent of clonal spread varying among species and forest stands. Although we did not find a clear pattern between clonality and forest age, we observed a greater number of clonal patches in secondary over primary forest stands.

Succession and seasonality of a Brazilian secondary tropical dry forest: Phenology and climate moderation

With large tracts of Tropical Dry Forests (TDF) in the Americas previously deforested, many places are currently undergoing secondary ecological succession. These processes vary in time and space, creating a landscape with forests of different ages or successional stages. These secondary forests need moreneed more understanding about how their ecosystem functionality is restored and recovering along the succession path. In particular, it is poorly understood how different successional stages moderate extremes of Essential Climatic Variables (ECV) such as temperature, relative humidity, soil moisture, and the fraction of Absorbed Photosynthetic Active Radiation (fAPAR). This paper, therefore, studies the linkages between high temporal resolution tower-based optical canopy phenology time series and seasonal changes in local ECV conditions between early (10–12 years) and late (50+ years) successional stage TDF in southeastern Brazil. We compare diurnal and seasonal patterns of understory ECVs between early/late plots and investigate the canopy’s Climate Moderating Capacity (CMC). Our data indicate at least 100 mm of precipitation within two weeks is necessary to initiate leaf flushing in this TDF. We also find that soil moisture and daytime air humidity strongly correlate (R2 > 0.75) with seasonal patterns in canopy vegetation greenness indices and fAPAR with a cross-correlation time lag peak of 12–15 days. Our results suggest that the late stage is more effective at cooling (−1.4 ± 0.6 °C vs −0.8 ± 0.6 °C) and reducing daytime evaporation (−0.41 ± 0.2 kPa vs −0.18 ± 0.12 °C) in the understory during the growing season than the early stage. Results suggest that the CMC varies with seasonal canopy leaf area dynamics and with heat and moisture scales, becoming significantly stronger when temperatures and vapour pressure deficits were highest, and correlated strongly with canopy fAPAR dynamics (R2 = 0.82). These findings demonstrate that TDFs can effectively regain microclimatic buffering capacity.

Seeding success: Integrating seed dispersal networks in tropical forest restoration

AbstractWhile the reassembly of fruit‐frugivore interactions remains at the forefront of tropical forest restoration, seed dispersal networks emerge as a potential approach to enhance restoration success. This review explores the integration of seed dispersal networks in tropical forest restoration, with the aims of (1) synthesizing important findings in the literature, (2) detailing potential biases in utilizing network theory, and (3) addressing current knowledge gaps and future directions for the field. We first highlight the importance of combining phytocentric and zoocentric approaches when sampling for seed dispersal interactions, as different methodologies have varying effects on network measures, and combining approaches can foster a more comprehensive understanding of dispersal interactions. Furthermore, when integrating seed dispersal networks into restoration goals, we suggest a highly connected and species‐rich network is desirable for earlier stages of forest succession where community turnover and transient interactions are pivotal. Nested patterns may emerge throughout varying stages of forest succession, and identifying generalist species that make up nested patterns may be useful for restoration practitioners in both early and later stages of forest regeneration. Modularity should be highest at later stages of succession to maintain community structure and stability, and connector species may play important roles in facilitating seed dispersal across temporal scales. Finally, we emphasize the importance of site‐specific long‐term datasets, chronosequences, and studies at large spatial scales to continue to understand network reassembly as a function of tropical forest succession and to develop effective strategies that enhance the recovery of tropical forest ecosystems.

Tropical forest succession increases tree taxonomic and functional richness but decreases evenness

AbstractAimSuccessional changes in functional diversity provide insights into community assembly by indicating how species are filtered into local communities based on their traits. Here, we assess successional changes in taxonomic and functional richness, evenness and redundancy along gradients of climate, soil pH and forest cover.LocationNeotropics.Time periodLast 0–100 years.Major taxa studiedTrees.MethodsWe used 22 forest chronosequence studies and 676 plots across the Neotropics to analyse successional changes in Hill's taxonomic and functional diversity of trees, and how these successional changes vary with continental‐scale gradients in precipitation, soil pH and surrounding forest cover.ResultsTaxonomic and functional richness and functional redundancy increased, while taxonomic and functional evenness decreased over time. Functional richness and evenness changed strongly when not accounting for taxonomic richness, but changed more weakly after statistically accounting for taxonomic richness, indicating that changes in functional diversity are largely driven by taxonomic richness. Nevertheless, the successional increases in functional richness when correcting for taxonomic richness may indicate that environmental heterogeneity and limiting similarity increase during succession. The taxonomically‐independent successional decreases in functional evenness may indicate that stronger filtering and competition select for dominant species with similar trait values, while many rare species and traits are added to the community. Such filtering and competition may also lead to increased functional redundancy. The changes in taxonomically‐independent functional diversity varied with resource availability and were stronger in harsh, resource‐poor environments, but weak in benign, productive environments. Hence, in resource‐poor environments, environmental filtering and facilitation are important, whereas in productive environments, weaker abiotic filtering allows for high initial functional diversity and weak successional changes.Main conclusionWe found that taxonomic and functional richness and functional redundancy increased and taxonomic and functional evenness decreased during succession, mainly caused by the increasing number of rare species and traits due to the arrival of new species and due to changing (a)biotic filters.

Trees adjust nutrient acquisition strategies across tropical forest secondary succession.

Nutrient limitation may constrain the ability of recovering and mature tropical forests to serve as a carbon sink. However, it is unclear to what extent trees can utilize nutrient acquisition strategies - especially root phosphatase enzymes and mycorrhizal symbioses - to overcome low nutrient availability across secondary succession. Using a large-scale, full factorial nitrogen and phosphorus fertilization experiment of 76 plots along a secondary successional gradient in lowland wet tropical forests of Panama, we tested the extent to which root phosphatase enzyme activity and mycorrhizal colonization are flexible, and if investment shifts over succession, reflective of changing nutrient limitation. We also conducted a meta-analysis to test how tropical trees adjust these strategies in response to nutrient additions and across succession. We find that tropical trees are dynamic, adjusting investment in strategies - particularly root phosphatase - in response to changing nutrient conditions through succession. These changes reflect a shift from strong nitrogen to weak phosphorus limitation over succession. Our meta-analysis findings were consistent with our field study; we found more predictable responses of root phosphatase than mycorrhizal colonization to nutrient availability. Our findings suggest that nutrient acquisition strategies respond to nutrient availability and demand in tropical forests, likely critical for alleviating nutrient limitation.

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.

Crown cover of a dominant pioneer legume affects tree species regeneration in a secondary tropical dry forest

Background: Individual pioneer tree species often dominate early tropical dry forest succession and thereby affect possible successional pathways. Mimosa acantholoba var. eurycarpa is a highly dominant species in the tropical dry forest in Mexico.&#x0D; Hypothesis: Mimosa acantholoba var. eurycarpa exerts an inhibitory effect on the germination, establishment, survival, and early growth of light-demanding pioneers, while facilitating these processes for shade-tolerant old-growth forests species.&#x0D; Studied species: Lonchocarpus torresiorum, Lysiloma divaricatum, Mimosa acantholoba var. eurycarpa and Vachellia farnesiana.&#x0D; Study site and dates: Nizanda, Oaxaca, Mexico, 2020-2021.&#x0D; Methods: In 12 early successional plots, we applied three levels of crown cover removal (100, 50, and 0 %) of established trees of the dominant pioneer legume (M. acantholoba var. eurycarpa). We sowed seeds of the four study species in each experimental plot and recorded their germination, establishment, survival, and early growth over a 2-mo period.&#x0D; Results: The removal of crown cover of established M. acantholoba var. eurycarpa trees did not significantly affect germination. Lysiloma divaricatum had the highest germination probability, the fastest germination, and the highest establishment probability regardless of treatment. Lonchocarpus torresiorum had the highest establishment probability in plots where the crown cover of established M. acantholoba var. eurycarpa trees was removed The survival probability of both shade-tolerant species was highest in the 100 % removal treatment.&#x0D; Conclusions: Despite successful germination of shade-tolerant species, their survival is inhibited under the dense canopy of the dominant legume. Therefore, interventions to reduce the crown cover area of this dominant legume may stimulate forest recovery.

A flash in the pan? The population dynamics of a dominant pioneer species in tropical dry forest succession

AbstractGlobally, around half of all tropical forests are secondary communities which are recovering from previous disturbances. In these communities, dominant pioneers play a critical role in the successional dynamics due their ability to modify the environment, and thus to facilitate or hinder the performance of other species. In this study, we examined the population dynamics of the dominant pioneer species, Mimosa acantholoba var. eurycarpa, in a tropical dry forest during the process of secondary succession. We collected data from permanent plots over a period of 13 years and used integral projection models to analyze the vital rates and changes in population size. Most vital rates were negatively related to successional age, with resprouting, survival and recruitment of individuals responding more strongly, and individual growth rate more weakly. These vital‐rate trends translated into significant variation of population size over succession, with a maximum size at year four and a rapid decline thereafter. Overall, our findings suggest that this species benefits from its ability to display a large number of resprouts in very early stages of succession, contributing to the rapid increase in population density. However, as succession unfolds, a lack of resprouting and a decline in survival reduce its population size, to the point of local extinction. These results highlight the importance of resprouting in the early dynamics of tropical dry forest communities recovering from disturbance.

Land use legacies affect early tropical forest succession in Mexico

AbstractQuestionsAgricultural expansion is one of the dominant drivers of forest and biodiversity loss, and shifting cultivation is the most widely used form of agriculture in many tropical forest regions. Where forests have been cleared, they have the potential to recover once the land is abandoned. However, legacies of land use are often overlooked in successional studies, and a deeper understanding of this legacy effect is needed to define efficient restoration practices using natural or assisted regeneration. Here, we analysed how land‐use history affects soil properties and early succession on abandoned agricultural fields in two contrasting Mexican socio‐ecological systems.LocationMexico, Oaxaca and Chiapas.MethodsWe sampled soil and monitored vegetation for 2 years after agricultural abandonment, and interviewed landowners about their land‐use practices.ResultsLand‐use practices were clearly influenced by landowners’ social context (residence time, rural or urban origin), and topography and soil type also constrained or facilitated land‐use practices. Soil characteristics were strongly affected by three land‐use practices: mechanical tillage decreased soil N and K; frequent herbicide and pesticide use increased N and K; and for pasture systems, stocking density increased soil bulk density and decreased pH and N. High‐intensity land management practices, specifically use of machinery, had the highest impact on early forest succession. When machinery was not used, the frequency of land‐use practices, particularly weeding frequency, is the main factor influencing tree cover and sapling diversity.ConclusionsTo facilitate post‐agricultural forest recovery, we recommend restoration efforts using natural regeneration in areas with low previous land‐use intensity and frequency.

Diversity patterns along ecological succession in tropical dry forests: a multi‐taxonomic approach

Evaluating the diversity of multiple taxa is fundamental to understand community assembly and to assess the integrity and functionality of tropical secondary forests. In this study, we analyzed the natural regeneration of tropical dry forests (TDFs) in three regions of Brazil using Hill–Simpson diversity, abundance and β‐diversity of trees and five groups of insects (herbivores, fruit‐feeding butterflies, ants, culicid mosquitoes and dung beetles). Sampling was conducted in 39 0.1 ha plots using a chronosequence approach (13 plots in early, intermediate and old‐growth forests). We evaluated the contribution of three different levels to γ‐diversity: α (within plots), β1 (among plots) and β2 (among successional stages), and further determined the relative importance of turnover (species replacement) and nestedness (differences in species number among sites) to β2. Our results showed that, unexpectedly, the Hill–Simpson diversity was consistently higher in early than old‐growth stages for all regions, but varied more widely in the intermediate stages. For each group separately, the same trend was observed for butterflies, ants, dung beetles and herbivores and did not differ among stages for mosquitoes. Successional differences in abundance were only detected for trees (increasing along the gradient) and for mosquitoes (decreasing). According to our expectations, the additive partitioning analysis showed that β2‐diversity contributed more to γ‐diversity than β1‐diversity, when all taxa were considered together and for most of them separately (except for butterflies and dung beetles). Most of the β2‐diversity was due to species turnover, but this contribution varied among groups and regions, with the highest turnover for herbivores and the lowest for dung beetles. Our results suggest that the Hill–Simpson diversity and changes in species composition (as given by β2‐diversity) are better indicators of forest natural regeneration than raw species richness, corroborating previous studies with plants and animals.

Biomass recovery along a tropical forest succession: Trends on tree diversity, wood traits and stand structure

Secondary forests recovering after disturbances currently comprise about half of the world′s tropical forests. A better understanding of the recovery of species composition, stand structure, and biomass stocks of these secondary forests is critical for the sustainable management of these ecosystems. The main objective of this research was to develop the successional trends in forest biomass, tree diversity, and species composition recovery. We monitored the changes in tree species composition, stand structural characteristics, and biomass growth in a chronosequence of tropical secondary forests in the Yucatan Peninsula of Southeastern Mexico. We used both linear and non-linear regressions to develop biomass growth models during forest succession. We calculated tree species diversity and similitude indices between forest successional groups (young, medium-aged, and old-growth forests). The aboveground biomass stocks ranged from 5.2 to 121 Mg C ha-1, which increased gradually with forest age. We observed a divergent trend of biomass accumulation between soft- and densewood species during succession. The contribution of densewood species on tree biomass stock increased (from 32 % to 81 %) with the increasing forest ages. However, the tree diversity indices did not change significantly with forest age and were a poor predictor of aboveground biomass growth during succession. Secondary forests (SFs) of advanced age were more similar to old-growth forests (OF) in their species composition compared to young ones, as evidenced by the Chao-Jaccard index. However, it may take more than a century for these secondary forests to recover to OF-level species composition. Furthermore, the structural equation model allowed us to infer the relative importance of forest age, tree diversity, soil properties, and stand structural attributes, including wood density on biomass growth along the forest chronosequence. Forest age-mediated effect of species composition shift and stand structural changes was a better predictor of biomass accumulation than the changes in tree diversity or soil properties during succession. However, the soil properties as a construct (latent variable), had a significant positive covariance with stand structure, indirectly influencing biomass growth. These biomass and species composition recovery models could be useful in forest management for biodiversity conservation, landscape restoration, and ecosystem services, including carbon sequestration.

Cost‐effectiveness of management strategies in a nucleation experiment in a tropical dry forest

Cost‐effective, large‐scale strategies are needed to restore degraded ecosystems worldwide. Applied nucleation is one technique that can accelerate succession in tropical forests. However, the effectiveness of irrigation and fertilization in the context of large‐scale applied nucleation in tropical dry forests (TDFs) has not yet been tested. To this end, we established a large‐scale experiment in southwestern Colombia on abandoned pastures. We planted 11,382 seedlings of 11 native species coupled with six management treatments that varied in the amounts of fertilizer and water. We monitored survival, height, and resprouting ability seedlings over 2 years. We also estimated the cost of seedling production, planting procedures, management, and monitoring, and assessed their cost‐effectiveness using seedling survival as an indication of effectiveness. After 2 years, 73% (8,266) of seedlings planted survived, and species survival ranged from 5 to 99%. Individuals that received the lowest amount of fertilizer (25 g of Nitrogen, Phosphorus, and Potassium [NPK]) with additional irrigation had the highest survival rates (&gt;76%). Final height ranged from 52 to 330 cm across species. Seedlings that received the highest amount of fertilizer (50 g of NPK) without additional irrigation had the highest mean final height (174 ± 9.0 cm). The control was cheapest ($7313/ha) and the most cost‐effective method in terms of seedling survival. Our findings suggest that the best way to establish cost‐effective, large‐scale restoration projects in TDFs is to plant native species mixtures of locally adapted species without fertilization and irrigation and to invest in long‐term monitoring.

The Changes, Aggregation Processes, and Driving Factors for Soil Fungal Communities during Tropical Forest Restoration.

Soil fungal communities play crucial roles in mediating the functional associations between above- and belowground components during forest restoration. Forest restoration shapes the alterations in plant and soil environments, which exerts a crucial effect on soil fungal assemblages. However, the changes, assembly processes, and driving factors of soil fungi communities during tropical forest restoration are still uncertain. We used Illumina high-throughput sequencing to identify the changes of soil fungal communities across a tropical secondary forest succession chronosequence (i.e., 12-, 42-, and 53-yr stages) in Xishuangbanna. During forest restoration, the dominant taxa of soil fungi communities shifted from r- to K-strategists. The relative abundance of Ascomycota (r-strategists) decreased by 10.0% and that of Basidiomycota (K-strategists) increased by 4.9% at the 53-yr restoration stage compared with the 12-yr stage. From the 12-yr to 53-yr stage, the operational taxonomic unit (OTU), abundance-based coverage estimator (ACE), Chao1, and Shannon index of fungal communities declined by 14.5-57.4%. Although the stochastic processes were relatively important in determining fungal assemblages at the late stage, the fungal community assembly was dominated by deterministic processes rather than stochastic processes. The shifts in soil properties resulting from tropical forest restoration exerted significant effects on fungal composition and diversity. The positive effects of microbial biomass carbon, readily oxidizable carbon, and soil water content explained 11.5%, 9.6%, and 9.1% of the variations in fungal community composition, respectively. In contrast, microbial biomass carbon (40.0%), readily oxidizable carbon (14.0%), and total nitrogen (13.6%) negatively contributed to the variations in fungal community diversity. Our data suggested that the changes in fungal composition and diversity during tropical forest restoration were primarily mediated by the positive or negative impacts of soil carbon and nitrogen pools.

Herbaceous species and dry forest species have more acquisitive leaf traits than woody species and wet forest species

Abstract Life forms are key to understand the changes in species composition and vegetation types in space and with succession over time. We asked how life forms from dry and wet regions differ in their leaf traits and trait hyperspace (i.e. trait variation) in early tropical forest succession on abandoned agricultural fields. We compared 324 early successional species from six life forms (herbs, grasses, vines, lianas, shrubs and trees) that occur in the first 5 years of succession on abandoned fields in Ghanaian wet and drought deciduous tropical forests. We measured 12 leaf traits that are important for carbon, water and nutrient use. A principal component analysis showed that 46% of trait variation is captured by a two‐dimensional spectrum of plant form and function: a leaf economics spectrum underpinning fast‐slow growth strategies, and a leaf size spectrum related to plant size and heat balance, which underlie species sorting along environmental gradients in space and time (during succession). Herbaceous and woody life forms had different leaf economics strategies: herbaceous species had more acquisitive trait values (e.g. higher leaf nutrient concentrations) that increase resource capture and use efficiency whereas woody species had more conservative trait values (e.g. higher leaf mass per area) that increase resource conservation. Regardless of life forms, dry forest species and deciduous species had more acquisitive trait values than wet forest species and evergreen species as they maximize their growth during the shorter growing season. The trait hyperspace was larger for woody life forms especially in wet forests. Synthesis. Herbaceous species had ‘faster’ leaf economics trait values and, hence, rapid carbon gain, explaining their success earlier in succession. In contrast, woody species had ‘slower’ resource conservation trait values that increase persistence, thus explaining their success later in succession. Deciduous species especially in a dry region had higher leaf nitrogen and phosphorus concentration to attain faster carbon gain to cope with a seasonally harsh environment. The trait hyperspace was generally smaller in the dry forest, as there are more deciduous species, and as a result, there is less variation in leaf lifespan and associated traits. Read the free Plain Language Summary for this article on the Journal blog.

Defaunated and invaded insular tropical rainforests will not recover alone: Recruitment limitation factors disentangled by hierarchical models of spontaneous and assisted regeneration

Abstract Most tropical forests are now severely degraded and their ability to recover is highly dependent on frugivores which ensure seed dispersal for most woody plants. The global collapse of large vertebrates therefore raises major concerns about tropical forest succession, but few field studies have been conducted to disentangle recruitment limitations during disrupted succession. This study took place on Réunion (Mascarenes) where all large native frugivores have been extinct since human colonisation in 1665 and where multiple invasions threaten native ecosystems. We set up 20 experimental blocks on a lava flow dated back to 1800, in plant‐impoverished post‐defaunation vegetation bordered by old‐growth forests. We assessed fecundity, seed dispersal and seedling recruitment of the complete fleshy‐fruited plant community and used Bayesian analyses to disentangle the impact of multiple factors on these key processes. In the same blocks, we sowed four native trees assumed to be disperserless to test their capacity to establish, controlling for two additional post‐dispersal limitations (seed predation and competition with invasive plants). On the flow, small‐seeded native plants were fairly dispersed but did not recruit, probably due to strong competition with invasive plants; the few native species that recruited somehow were mostly medium‐seeded plants that were still dispersed; large‐seeded plants were absent from seed rain (which shows that invasive frugivores did not replace extinct ones) and subsequently from spontaneous recruitment. Instead, some alien plants, notably the tiny‐seeded highly dispersed Clidemia hirta and the medium‐seeded Psidium cattleianum largely dominated seedling recruitment. Native plants recruited better at the forest margin, including some large‐seeded species nearby mother trees. Sown large‐seeded species were able to emerge and survive in all plots whatever the treatment, which demonstrates that dispersal loss was the primary cause of regeneration failure on the flow. Synthesis. The strong modulation of the establishment capacity of native plants by seed mass shows that invasive plants win by forfeit of large‐seeded plants after native frugivores loss. Our study emphasises the fundamental role of dispersal loss and competition with invasive plants in the disruption of ecological succession, as well as the urgency of restoring seed dispersal and strengthening biosecurity regulations.

Effects of large mammal exclusion on seedling communities depend on plant species traits and landscape protection in human‐modified Costa Rican forests

Abstract Large terrestrial herbivorous mammals (LTH‐mammals) influence plant community structure by affecting seedling establishment in mature tropical forests. Many of these LTH‐mammals frequent secondary forests, but their effects on seedling establishment in them are understudied, hindering our understanding of how LTH‐mammals influence forest regeneration in human‐modified landscapes. We tested the hypothesis that the strength of LTH‐mammals' effects on seedling establishment depends on landscape protection, forest successional stage and plant species' traits using a manipulative field experiment in six 1‐ha sites with varying successional age and landscape protection. In each site, we established 40 seedling plot‐pairs, with one plot excluding LTH‐mammals and one not, and monitored seedlings of 116 woody species for 26 months. We found significant effects of LTH‐mammal exclusion on seedling survival contingent upon the protection of forests at the landscape level and forest stage. After 26 months, survival differences between LTH‐mammal exclusion and non‐exclusion treatments were greater in protected than unprotected landscapes. Additionally, plant species' traits were related to the LTH‐mammals' differential effects, as LTH‐mammals reduced the survival of seedlings of larger‐seeded species the most. Overall, LTH‐mammals' effects translated into significant shifts in community composition as seedling communities inside and outside the exclosures diverged. Moreover, lower density and higher species diversity were found as early as 12 and 18 months outside than inside exclosures. Synthesis and applications. Insight into the interactions between LTH‐mammals and seedling communities in forest regeneration can be instrumental in planning effective restoration efforts. We highlight the importance of landscape protection in seedling survival and the role of LTH‐mammals in promoting seedling diversity in mature forests but also in secondary successional forests. The findings suggest that conservation efforts and possibly trophic rewilding can be important approaches for preserving diversity and influencing the trajectory of secondary tropical forest succession. However, we also caution that an overabundance of LTH‐mammals may adversely impact the pace of forest succession due to their preference for large‐seeded species. Therefore, a comprehensive wildlife management plan is indispensable. Additionally, longer term studies on LTH‐mammals are necessary to understand the effects of temporal fluctuations that are undetected in short‐term studies.

Incomplete recovery of tree community composition and rare species after 120 years of tropical forest succession in Panama.

Determining how fully tropical forests regenerating on abandoned land recover characteristics of old-growth forests is increasingly important for understanding their role in conserving rare species and maintaining ecosystem services. Despite this, our understanding of forest structure and community composition recovery throughout succession is incomplete, as many tropical chronosequences do not extend beyond the first 50 years of succession. Here, we examined trajectories of forest recovery across eight 1-hectare plots in middle and later stages of forest succession (40-120 years) and five 1-hectare old-growth plots, in the Barro Colorado Nature Monument (BCNM), Panama. We first verified that forest age had a greater effect than edaphic or topographic variation on forest structure, diversity and composition and then corroborated results from smaller plots censused 20 years previously. Tree species diversity (but not species richness) and forest structure had fully recovered to old-growth levels by 40 and 90 years, respectively. However, rare species were missing, and old-growth specialists were in low abundance, in the mid- and late secondary forest plots, leading to incomplete recovery of species composition even by 120 years into succession. We also found evidence that dominance early in succession by a long-lived pioneer led to altered forest structure and delayed recovery of species diversity and composition well past a century after land abandonment. Our results illustrate the critical importance of old-growth and old secondary forests for biodiversity conservation, given that recovery of community composition may take several centuries, particularly when a long-lived pioneer dominates in early succession. Abstract in Spanish is available with online material.

Shifting cultivation and logging change soil organic carbon functional groups in tropical lowland rainforests on Hainan Island in China

Recent evidence has demonstrated that tropical forest secondary succession has significant influences on soil organic C (SOC) storage. However, the changes in the SOC functional groups with forest restoration are still uncertain. In this study, the stock and functional groups of SOC in tropical forest recovery after two types of disturbance were measured, and the key influencing factors were analyzed. The SOC stocks were significantly higher in the young-secondary forests after shifting cultivation and in the clear cut forests than in the old-growth forests. However, the shifting cultivation and logging practices significantly increased the ratio of carbohydrate C to N-alkyl/methoxyl C (CC/MC), indicating that the decomposition degree of SOC in the old-growth forests was higher. The proportion of aromatic C in soil was higher in the young-secondary forests after shifting cultivation than in the old-growth forests. The proportion of aromatic C in litter and fine roots, the α-diversity of soil fungal and bacterial communities, and soil phosphorus were positively correlated with soil aromatic C between the shifting cultivation secondary and old-growth forests. The proportion of O-alkyl C in soil was higher in the logged forests than in the old-growth forests. The proportion of O-alkyl C in litter was positively correlated with O-alkyl C in soil between the logged and old-growth forests. Our results suggest that the labile O-alkyl C functional group recovers more rapidly after logging practices, and restoration after shifting cultivation increases the labile carbohydrate C functional group and aromaticity in tropical lowland rainforests. However, SOC has greater chemical stability in terms of the decomposition degree in tropical old-growth forests.