Leaf litter arthropod responses to tropical forest restoration

Soil and litter arthropods represent a large proportion of tropical biodiversity and perform important ecosystem functions, but little is known about the efficacy of different tropical forest restoration strategies in facilitating their recovery in degraded habitats. We sampled arthropods in four 7‐ to 8‐year‐old restoration treatments and in nearby reference forests. Sampling was conducted during the wet and dry seasons using extractions from litter and pitfall samples. Restoration treatments were replicated in 50 × 50‐m plots in four former pasture sites in southern Costa Rica: plantation – trees planted throughout the plot; applied nucleation/islands – trees planted in patches of different sizes; and natural regeneration – no tree planting. Arthropod abundance, measures of richness and diversity, and a number of functional groups were greater in the island treatment than in natural regeneration or plantation treatments and, in many cases, were similar to reference forest. Litter and pitfall morphospecies and functional group composition in all three restoration treatments were significantly different than reference sites, but island and plantation treatments showed more recovery than natural regeneration. Abundance and functional group diversity showed a much greater degree of recovery than community composition. Synthesis and applications: The less resource‐intensive restoration strategy of planting tree islands was more effective than tree plantations in restoring arthropod abundance, richness, and functional diversity. None of the restoration strategies, however, resulted in similar community composition as reference forest after 8 years of recovery, highlighting the slow rate of recovery of arthropod communities after disturbance, and underscoring the importance of conservation of remnant forests in fragmented landscapes.     

Recovery of floristic diversity and basal area in natural forest regeneration and planted plots in a Costa Rican wet forest

When compared to planted reforestation, natural unassisted regeneration is often reported to result in slow recovery of biomass and biodiversity, especially early in succession. In some cases, naturally regenerating forests are not comparable to the community structure of primary forests after many decades. However, direct comparison of the outcomes of tropical forest restoration and natural regeneration is hindered by differences in metrics of forest recovery, inconsistency in land use histories, and dissimilarities in experimental design. We present the results of a replicated reforestation experiment comparing natural regeneration and polyculture tree planting at multiple diversity levels (3, 6, 9, or 12 native tree species), with uniform land use history and initial edaphic conditions. We compare the recovery of basal area and floristic diversity in these treatments after 5 yr of succession. Total basal area was higher in planted plots than in naturally regenerating plots, but it but did not vary among the different planted diversity levels. The basal area of woody recruits did not differ among treatments. The diversity of woody recruits increased substantially over time but did not vary among planting treatments. Species composition trajectories showed directional turnover over time, with no consistent differences among treatments. The convergence of restoration trajectories and similarity of floristic community diversity and composition across all treatments, after only 5 yr, provides evidence of the viability of natural regeneration for rapid restoration of forest biodiversity.     

Small Clusters of Fast-Growing Trees Enhance Forest Structure on Restored Bottomland Sites

Despite the diversity of trees in bottomland forests, restoration on bottomland sites is often initiated by planting only a few species of slow‐growing, hard mast–producing trees. Although successful at establishing trees, these young forests are slow to develop vertical structure, which is a key predictor of forest bird colonization. Furthermore, when natural seed sources are few, restored sites may be depauperate in woody species. To increase richness of woody species, maximum tree height, and total stem density, I supplemented traditional plantings on each of 40 bottomland restoration sites by planting 96 Eastern cottonwood (Populus deltoides) and American sycamore (Platanus occidentalis) in eight clusters of 12 trees. First‐year survival of cottonwood stem cuttings (25%) and sycamore seedlings (47%) was poor, but survival increased when afforded protection from competition with weeds. After five growing seasons, 165 of these 320 supplemental tree clusters had at least one surviving tree. Vegetation surrounding surviving clusters of supplemental trees harbored a greater number of woody species, increased stem density, and greater maximum tree height than was found on paired restoration sites without supplemental trees. These increases were primarily accounted for by the supplemental trees.     

Scale‐dependent effects of forest restoration on Neotropical fruit bats

Neotropical fruit bats (family Phyllostomidae) facilitate forest regeneration on degraded lands by dispersing shrub and tree seeds. Accordingly, if fruit bats can be attracted to restoration sites, seed dispersal could be enhanced. We surveyed bat communities at 10 sites in southern Costa Rica to evaluate whether restoration treatments attracted more fruit bats if trees were planted on degraded farmlands in plantations or island configurations versus natural regeneration. We also compared the relative influence of tree cover at local and landscape spatial scales on bat abundances. We captured 68% more fruit bat individuals in tree plantations as in controls, whereas tree island plots were intermediate. Bat activity also responded to landscape tree cover within a 200‐m radius of restoration plots, with greater abundance but lower species richness in deforested landscapes. Fruit bat captures in controls and tree island plots declined with increasing landscape tree cover, but captures in plantations were relatively constant. Individual species responded differentially to tree cover measured at different spatial scales. We attribute restoration effects primarily to habitat structure rather than food resources because no planted trees produced fruits regularly eaten by bats. The magnitude of tree planting effects on fruit bats was less than previous studies have found for frugivorous birds, suggesting that bats may play a particularly important role in dispersing seeds in heavily deforested and naturally regenerating areas. Nonetheless, our results show that larger tree plantations in more intact landscapes are more likely to attract diverse fruit bats, potentially enhancing seed dispersal.     

Application of Assisted Natural Regeneration to Restore Degraded Tropical Forestlands

Assisted natural regeneration (ANR) is a simple, low‐cost forest restoration method that can effectively convert deforested lands of degraded vegetation to more productive forests. The method aims to accelerate, rather than replace, natural successional processes by removing or reducing barriers to natural forest regeneration such as soil degradation, competition with weedy species, and recurring disturbances (e.g., fire, grazing, and wood harvesting). Compared to conventional reforestation methods involving planting of tree seedlings, ANR offers significant cost advantages because it reduces or eliminates the costs associated with propagating, raising, and planting seedlings. It is most effectively utilized at the landscape level in restoring the protective functions of forests such as watershed protection and soil conservation. ANR techniques are flexible and allow for the integration of various values such as timber production, biodiversity recovery, and cultivation of crops, fruit trees, and non‐timber forest products in the restored forest. This paper describes the steps of applying ANR and conditions under which it will be most effective. It also discusses ANR’s comparative advantages as well as some of its constraints.     

Natural Regeneration in Plantations of Native Trees in Lowland Brazilian Atlantic Forest: Community Structure,Diversity, and Dispersal Syndromes

Plantations of native‐tree species are often recommended for ecological restoration, but the understanding of how these techniques catalyze natural ecological processes is limited. We investigated natural regeneration in five plantations of native trees in the Poço das Antas Biological Reserve (PABR) in the state of Rio de Janeiro, Brazil. The plantations were 9–11 years old, and contained 8–14 native‐tree species with different compositions and relative density of species. We analyzed floristic composition, structure (density and basal area) of overstory and understory strata, as well as other ecological attributes (dispersal syndromes, fruit or seed size, and the availability of fruit for frugivores). Zoochorous species comprised 77% of the community, with a prevalence of the two smallest size classes of propagules (< 0.6 and 0.6–1.6 cm) in natural regeneration. The density of zoochorous plants in the understory was positively correlated with their density in the overstory, indicating their influence on natural regeneration (r² = 0.36; p < 0.0002). Fruit availability for frugivores (density and richness of plants fruiting during the year) was also positively correlated with the density of stems in the understory. Therefore, attributes such as dispersal syndrome and fruiting season should be considered in selecting species to be planted. The differences in natural regeneration observed in each of the native‐tree plantations indicated that the performance of plantations as a restoration strategy may differ, depending on initial species composition, planting density, and site conditions.     

Ecological outcomes of agroforests and restoration 15 years after planting

Large‐scale forest restoration relies on approaches that are cost‐effective and economically attractive to farmers, and in this context agroforestry systems may be a valuable option. Here, we compared ecological outcomes among (1) 12–15‐year‐old coffee agroforests established with several native shade trees, (2) 12–15‐year‐old high‐diversity restoration plantations, and (3) reference old‐growth forests, within a landscape restoration project in the Pontal do Paranapanema region, in the Atlantic Forest of southeastern Brazil. We compared the aboveground biomass, canopy cover, and abundance, richness, and composition of trees, and the regenerating saplings in the three forest types. In addition, we investigated the landscape drivers of natural regeneration in the restoration plantations and coffee agroforests. Reference forests had a higher abundance of trees and regenerating saplings, but had similar levels of species richness compared to coffee agroforests. High‐diversity agroforests and restoration plantations did not differ in tree abundance. However, compared to restoration plantations, agroforests showed higher abundance and species richness of regenerating saplings, a higher proportion of animal‐dispersed species, and higher canopy cover. The abundance of regenerating saplings declined with increasing density of coffee plants, thus indicating a potential trade‐off between productivity and ecological benefits. High‐diversity coffee agroforests provide a cost‐effective and ecologically viable alternative to high‐diversity native tree plantations for large‐scale forest restoration within agricultural landscapes managed by local communities, and should be included as part of the portfolio of reforestation options used to promote the global agenda on forest and landscape restoration.     

Litterfall Dynamics Under Different Tropical Forest Restoration Strategies in Costa Rica

In degraded tropical pastures, active restoration strategies have the potential to facilitate forest regrowth at rates that are faster than natural recovery, enhancing litterfall, and nutrient inputs to the forest floor. We evaluated litterfall and nutrient dynamics under four treatments: plantation (entire area planted), tree islands (planting in six patches of three sizes), control (same age natural regeneration), and young secondary forest (7–9‐yr‐old natural regeneration). Treatments were established in plots of 50 × 50 m at six replicate sites in southern Costa Rica and the annual litterfall production was measured 5 yr after treatment establishment. Planted species included two native timber‐producing hardwoods (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N‐fixing species (Inga edulis and Erythrina poeppigiana). Litter production was highest in secondary forests (7.3 Mg/ha/yr) and plantations (6.3), intermediate in islands (3.5), and lowest in controls (1.4). Secondary forests had higher input of all nutrients except N when compared with the plantation plots. Inga contributed 70 percent of leaffall in the plantations, demonstrating the influence that one species can have on litter quantity and quality. Although tree islands had lower litterfall rates, they were similar to plantations in inputs of Mg, K, P, Zn, and Mn. Tree islands increased litter production and nutrient inputs more quickly than natural regeneration. In addition to being less resource intensive than conventional plantations, this planting design promotes a more rapid increase in litter diversity and more spatial heterogeneity, which can accelerate the rate of nutrient cycling and facilitate forest recovery.     

Native tree regeneration in native tree plantations: understanding the contribution of Araucaria angustifolia to biodiversity conservation in the threatened Atlantic Forest in Argentina

Deforestation is a global process that has strongly affected the Atlantic Forest in South America, which has been recognised as a threatened biodiversity hotspot. An important proportion of deforested areas were converted to forest plantations. Araucaria angustifolia is a native tree to the Atlantic Forest, which has been largely exploited for wood production and is currently cultivated in commercial plantations. An important question is to what extent such native tree plantations can be managed to reduce biodiversity loss in a highly diverse and vulnerable forest region . We evaluated the effect of stand age, stand basal area, as a measure of stand density, and time since last logging on the density and richness of native tree regeneration in planted araucaria stands that were successively logged over 60 years, as well as the differences between successional groups in the response of plant density to stand variables. We also compared native tree species richness in planted araucaria stands to neighbouring native forest. Species richness was 71 in the planted stands (27 ha sampled) and 82 in native forest (18 ha sampled) which approximate the range of variation in species richness found in the native forests of the study area. The total abundance and species richness of native trees increased with stand age and time since last logging, but ecological groups differed in their response to such variables. Early secondary trees increased in abundance with stand age 3–8 times faster than climax or late secondary trees. Thus, the change in species composition is expected to continue for a long term. The difference in species richness between native forest and planted stands might be mainly explained by the difference in plant density. Therefore, species richness in plantations can contribute to local native tree diversity if practices that increase native tree density are implemented.     

Enrichment planting does not improve tree restoration when compared with natural regeneration in a former pine plantation in Kibale National Park,Uganda

Given the high rates of deforestation and subsequent land abandonment, there are increasing calls to reforest degraded lands; however, many areas are in a state of arrested succession. Plantations can break arrested succession and the sale of timber can pay for restoration efforts. However, if the harvest damages native regeneration, it may be necessary to intervene with enrichment planting. Unfortunately, it is not clear when intervention is necessary. Here, we document the rate of biomass accumulation of planted seedlings relative to natural regeneration in a harvested plantation in Kibale National Park, Uganda. We established two 2‐ha plots and in one, we planted 100 seedlings of each of four native species, and we monitored all tree regeneration in this area and the control plot. After 4 years, naturally regenerating trees were much taller, larger and more common than the planted seedlings. Species richness and two nonparametric estimators of richness were comparable between the plots. The cumulative biomass of planted seedlings accounted for 0.04% of the total above‐ground tree biomass. The use of plantations facilitated the growth of indigenous trees, and enrichment planting subsequent to harvesting was not necessary to obtain a rich tree community with a large number of new recruits.     

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.     

Restoration of Tropical Moist Forests on Bauxite-Mined Lands in the Brazilian Amazon

We evaluated forest structure and composition in 9- to 13-year-old stands established on a bauxite-mined site at Trombetas (Pará), Brazil, using four different reforestation techniques following initial site preparation and topsoil replacement. These techniques included reliance on natural forest regeneration, mixed commercial species plantings of mostly exotic timber trees, direct seeding with mostly native early successional tree species, and mixed native species plantings of more than 70 tree species (the current operational restoration treatment at this site). Replicated fixed-radius plots in each treatment and in undisturbed primary forest were used to quantify the canopy and understory structure and the abundance and diversity of all vascular plant species. Treatment comparisons considered regeneration density, species richness and diversity for all floristic categories, and, for trees and shrubs, the relative contribution of initial planting and subsequent regeneration from soil seed banks and seed inputs from nearby primary forests. With the possible exception of the stands of mixed commercial species, which were superior to all others in terms of tree basal-area development but relatively poor in species richness, all treatments were structurally and floristically diverse, with a high probability of long-term restoration success. Of these, the mixed native species plantings appeared to be at least risk of arrested succession due to the dominance of a broader range of tree species of different successional stages or expected life spans. In all treatments, several locally important families of primary forest trees (Annonaceae, Chrysobalanaceae, Lauraceae, Palmae and Sapotaceae) were markedly underrepresented due to a combination of poor survival of initial plantings and limitations on seed dispersal from the surrounding primary forest.     

Planting Design Effects on Avian Seed Dispersers in a Tropical Forest Restoration Experiment

Seed dispersal often limits tropical forest regeneration and animals disperse most rainforest tree seeds. This presents two important questions for restoration ecologists: (1) which animals are common seed dispersers? and (2) which restoration techniques attract them? Fourteen restoration sites were planted with four tree species in three designs, (1) controls (no planting, natural regeneration) (2) islands (trees planted in small patches), and (3) plantations (trees planted continuously over a large patch). We sampled birds in November, February, and April 2007–2008 with mist nets, in February and July 2009 with observations, and in July 2008 with both techniques. We documented 30 seed species from fecal samples of captured birds. All identified seed species were early‐successional forms. Four tanager species, three thrushes, two saltators, two flycatchers, and one finch were categorized as common seed dispersers, based on their high likelihood of dispersing seeds. Common dispersers were generalist species with small gape widths (<15 mm). Common dispersers were captured significantly more often in plantations than controls in most seasons and more often in plantations than islands during one season. Common disperser observations were significantly greater in plantations than controls during two periods and in plantations compared with islands in one period. Results indicate that plantation‐style planting is the conservative strategy to maximize attractiveness to common dispersers in tropical restoration sites. Island planting is an alternative when resources are limited although disperser activity may be lower in some seasons than in plantations. Additional research should investigate how to attract large, forest‐associated dispersers.     

Comparing the recovery of richness,structure, and biomass in naturally regrowing and planted reforestation

The clearing of natural vegetation for agriculture has reduced the capacity of natural systems to provide ecosystem functions. Ecological restoration can restore desirable ecosystem functions, such as creating habitat for animal conservation and carbon sequestration as woody biomass. In order to maintain these beneficial ecosystem functions, restoration projects need to mature into self‐perpetuating communities. Here we compared the ecological attributes of two types of restoration, “active” tree plantings with “passive” natural forest regeneration (“natural regrowth”) to existing remnant vegetation in a cleared agricultural landscape. Specifically, we measured differences between forest categories in factors that may predict future restoration failure or ecosystem collapse: aboveground plant biomass and biomass accrual over time (for regrowing stands), plant density and size class distributions, and diversity of functional groups based on seed dispersal and growth strategy traits. We found that natural regrowth and planted forests were similar in many ecological characteristics, including biomass accrual. Despite this, planted stands contained fewer tree recruit and shrub individuals, which may be due to limited recruitment in plantings. If this continues, these forests may be at risk of collapsing into nonforest states after mature trees senesce. Lower shrub density and richness of mid‐story trees may lead to lower structural complexity in planting plots, and alongside lower richness of fleshy‐fruited plant species may reduce animal resources and animal use of the restored stand. In our study region, natural regrowth may result in restored woodland communities with greater conservation and carbon mitigation value.     

Flower and Fruit Availability along a Forest Restoration Gradient

Frugivores and pollinators are two functional groups of animals that help ensure gene flow of plants among sites in landscapes under restoration and to accelerate restoration processes. Resource availability is postulated to be a key factor to structure animal communities using restoration sites, but it remains poorly studied. We expected that diverse forests with many plant growth forms that have less‐seasonal phenological patterns will provide more resources for animals than forests with fewer plant growth forms and strongly seasonal phenological patterns. We studied forests where original plantings included high tree species diversity. We studied resource provision (richness and abundance of flowers and fruits) of all plant growth forms, in three restoration sites of different ages compared to a reference forest, investigating whether plant phenology changes with restoration process. We recorded phenological data for reproductive plant individuals (351 species) with monthly sampling over 2 years, and found that flower and fruit production have been recovered after one decade of restoration, indicating resource provision for fauna. Our data suggest that a wide range of plant growth forms provides resource complementarities to those of planted tree species. Different flower phenologies between trees and non‐trees seem to be more evident in a forest with high non‐tree species diversity. We recommend examples of ideal species for planting, both at the time of initial planting and post‐planting during enrichment. These management actions can minimize shortage and periods of resource scarcity for frugivorous and nectarivorous fauna, increasing probability of restoring ecological processes and sustainability in restoration sites.     

坝上地区不同年代退耕还林生境的草本层植物多样性及影响因子

退耕还林是坝上地区重要的生态修复工程,其生态效益日益受到广泛关注,但针对林下植物多样性保护功能的研究还较少,虽然草本层植被是森林生态系统服务功能的重要指标之一。为评估不同年代退耕还林生境的生物多样性,调查了20世纪70年代至21世纪初共4个年代退耕还林生境的草本层植物多样性,并与弃耕生境及自然林生境进行了对比。为进一步分析林下草本层植物多样性的影响因子,还分析了乔灌层植被、土壤理化性质与林下草本植物多样性的关系。结果显示,不同年代退耕还林的林下草本植物多样性未有显著差异,且均显著低于弃耕生境与自然林生境。草本层物种丰富度对土壤碱解氮含量、灌木层丰富度、灌木层密度具有显著的正响应,Simpson多样性主要与乔木层林木密度呈显著负相关。研究表明,不同年代退耕还林后的林下草本层植物多样性保护功能低于预期,而改善林分空间结构与建设模式可能利于提升其生物多样性保护功能。未来植被修复中需充分权衡生态系统不同服务功能之间的关系,因地制宜,宜林则林,宜草则草,重视自然恢复与多元化生态修复方式,以实现更好生态效益。     

Testing a facilitation model for ecosystem restoration: Does tree planting restore ground layer species in a grassy woodland?

Planning for the restoration of degraded ecosystems has a strong basis in facilitation successional theory, which, as applied in restoration practice, states that planting of structurally dominant tree species will assist the entry of other native species into a restored community. In Australia, tree planting has been widely applied in restoration of grassy woodland ecosystems. Trees have been postulated to reduce the cover and diversity of weed species, thus facilitating recolonization of native woodland species (indirect facilitation). The expected outcomes of this process include reduced species richness and abundance of exotic plant species and increased species richness and abundance/dominance of natives in areas beneath tree canopies, with these trends strengthening with time. To assess whether this was occurring, we carried out a comparative analysis of species assemblages found underneath and outside of planted tree canopies in sites replanted with juvenile canopy tree species 3–5 or 8–10 years previously. We sampled revegetated stands of Cumberland Plain Woodland, an endangered ecological community in Western Sydney, Australia. We found that neither the number nor abundance of native ground layer species beneath canopies increased as a result of trees being planted at sites of both ages. Where seed is limited, we predicted an increase in abundance of existing native species under planted tree canopies. On this point, the results were mixed and showed some natives with an increased abundance while others decreased. Exotic species richness showed the reverse of the expected pattern, being greater under tree canopies. These findings lend no support to the theory of indirect facilitation. We conclude that simple facilitation models may be inadequate to support planning of grassy woodland restoration and that those models incorporating successional time lags and restoration barriers are likely to be more informative about the development of communities initiated by tree planting.     

Roles of non‐native species in large‐scale regeneration of moist tropical forests on anthropogenic grassland

This paper presents a new synthesis of the role of native and non‐native species in diverse pathways and processes that influence forest regeneration on anthropogenic grassland in the moist tropics. Because of altered species composition, abiotic conditions and landscape habitat mosaics, together with human interventions, these successional pathways differ from those seen in pre‐clearing forests. However, representation of different functional life forms of plant (tree, vine, grass, herb and fern) and animal (frugivorous seed disperser, granivorous seed predator, seedling herbivore and carnivore) shows consistent global variation among areas of pasture, intact forest, and post‐grassland regrowth. Biotic webs of interaction involve complex indirect influences and feedbacks, which can account for wide observed variation in regeneration trajectories over time. Important processes include: limitation of tree establishment by dense grasses; recruitment and growth of pioneer pasture trees (shading grasses and facilitating bird‐assisted seed dispersal); and smothering of trees by vines. In these interactions, species’ functional roles are more important than their biogeographic origins. Case studies in eastern Australia show native rain forest plant species diversity in all life forms increasing over time when pioneer trees are non‐native (e.g., Cinnamomum camphora, Solanum mauritianum), concurrent with decreased grass and fern cover and increased abundance of trees and vine tangles. The global literature shows both native and non‐native species facilitating and inhibiting regeneration. However conservation goals are often targeted at removing non‐native species. Achieving large‐scale tropical forest restoration will require increased recognition of their multiple roles, and compromises about allocating resources to their removal.     

Restoration of semi-natural forest after clearcutting of conifer plantations in Japan

We reviewed recent studies on the restoration of semi-natural forests after clearcutting of conifer plantations with specific reference to the importance of setting restoration goals, general and specific factors influencing restoration, and the prediction and judgment of successful restoration. For the restoration of semi-natural forests following the clearcutting of conifer plantations, recovery levels and appropriate restoration methods cannot be considered separately from the aims of restoration to provide ecosystem services. Restoration needs to be based on the setting of goals that correspond to the type and degree of ecosystem services targeted. When we aim to restore vegetation dominated by late-seral canopy trees within as short a period after the clearcutting of conifer plantations as possible, advanced regeneration in the pre-logged plantation is the most important and reliable source of post-logging regeneration, just as it is in the recovery of disturbed non-plantation forests. Advanced regeneration is not always present in plantations. Seedling establishment by seed rain from adjacent natural forest patches after a disturbance can contribute to late-seral or canopy species; however, the range of the edge effect is generally short, and the degree of the edge effect depends on the natural forest type and is more pronounced in deciduous forests, such as those in cool-temperate climates, or in forests with a complex forest structure. Thus, advanced regeneration is a better predictor of short-term forest recovery than measures based on seed rain from adjacent seed sources. However, restoration achieved only through advanced regeneration can simplify the species composition, and the recolonization processes of gradual species accession from seeds should be considered in the long-term context of restoration.     

Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.