Secondary forests comprise an increasing area of the tropics and play an important role in global carbon cycling. We compare above-ground biomass accumulation of both planted and naturally regenerating trees, as well as C in the top soil layer, in three restoration treatments replicated at 14, six to eight year old restoration sites in southern Costa Rica. Restoration strategies include: control (no planting), planting tree islands, and conventional, mixed-species tree plantations. We evaluate the importance of past land-use, soil nutrients, understory cover, and surrounding forest cover in explaining variation in above-ground biomass accumulation (ABA) rate across sites. Total ABA and planted tree ABA rate were highest in plantations, intermediate in islands, and lowest in control treatments, whereas ABA rate of naturally regenerating trees did not differ across treatments. Most ABA in plantations (89%) and islands (70%) was due to growth of planted trees. Soil carbon did not change significantly over the time period of the study in any treatment. The majority of across-site variation in both total and planted tree ABA rate was explained by duration of prior pasture use. Tree growth in the first two years after planting explained approximately two-thirds of the variation in ABA rate after 6-8. years. Soil nutrient concentrations explained relatively little of the variation in planted or naturally recruiting ABA rate. Our results show that planting trees substantially increases biomass accumulation during the first several years of forest recovery in former agricultural lands and that past-land use has a strong effect on the rate of biomass accumulation. Planting tree islands is a cost-effective strategy for increasing ABA and creating more heterogeneous habitat conditions than tree plantations. We recommend small scale planting trials to quickly assess potential biomass accumulation and prioritize sites for ecosystem service payments for carbon sequestration. © 2014 Elsevier B.V.

Past studies have shown that tropical forest regeneration on degraded farmlands is initially limited by lack of seed dispersal, but few studies have tracked changes in abundance and composition of seed rain past the first few years after land abandonment. We measured seed rain for 12 months in 10 6-9-year-old restoration sites and five mature, reference forests in southern Costa Rica in order to learn (1) if seed rain limitation persists past the first few years of regeneration; (2) how restoration treatments influence seed community structure and composition; and (3) whether seed rain limitation is contingent on landscape context. Each restoration site contained three 0.25-ha treatment plots: (1) a naturally regenerating control, (2) tree islands, and (3) a mixed-species tree plantation. Sites spanned a deforestation gradient with 9-89% forest area within 500 m around the treatment plots. Contrary to previous studies, we found that tree seeds were abundant and ubiquitous across all treatment plots (585.1 ± 142.0 seeds · m(-2) · yr(-1) [mean ± SE]), indicating that lack of seed rain ceased to limit forest regeneration within the first decade of recovery. Pioneer trees and shrubs comprised the vast majority of seeds, but compositional differences between restoration sites and reference forests were driven by rarer, large-seeded species. Large, animal-dispersed tree seeds were more abundant in tree islands (4.6 ± 2.9 seeds · m(-2) · yr(-1)) and plantations (5.8 ± 3.0 seeds · m(-2) · yr(-1)) than control plots (0.2 ± 0.1 seeds · m(-2) · yr(-1)), contributing to greater tree species richness in actively restored plots. Planted tree species accounted for < 1% of seeds. We found little evidence for landscape forest cover effects on seed rain, consistent with previous studies. We conclude that seed rain limitation shifted from an initial, complete lack of tree seeds to a specific limitation on large-seeded, mature forest species over the first decade. Although total seed abundance was equal among restoration treatments, tree plantations and tree islands continued to diversify seed rain communities compared to naturally regenerating controls. Compositional differences between regenerating plots and mature forests suggest that large-seeded tree species are appropriate candidates for enrichment planting.

We agree with Prach and del Moral that passive recovery is often a desirable and effective restoration strategy. Passive and various active restoration approaches need to be weighed on a case-by-case basis and depend on the goals, relative rates of recovery desired, and various social and financial costs implicit in each option. That said, we stress that passive restoration has a unique set of challenges and costs, which we highlighted in our original article and briefly reiterate here.

Both dispersal- and niche-based factors can impose major barriers on tree establishment. Our understanding of how these factors interact to determine recruitment rates is based primarily on findings from mature tropical forests, despite the fact that a majority of tropical forests are now secondary. Consequently, factors influencing seed limitation and the seed-to-seedling transition (STS) in disturbed landscapes, and how those factors shift during succession, are not well understood. We used a 3.5-yr record of seed rain and seedling establishment to investigate factors influencing tree recruitment after a decade of recovery in a tropical wet forest restoration experiment in southern Costa Rica. We asked (1) how do a range of restoration treatments (natural regeneration, applied nucleation, plantation), canopy cover, and life-history traits influence the STS and (2) how do seed and establishment limitation (lack of seed arrival or lack of seedling recruitment, respectively) influence vegetation recovery within restoration treatments as compared to remnant forest? We did not observe any differences in STS rates across restoration treatments. However, STS rates were lowest in adjacent later successional remnant forests, where seed source availability did not highly limit seed arrival, underscoring that niche-based processes may increasingly limit recruitment as succession unfolds. Additionally, larger-seeded species had consistently higher STS rates across treatments and remnant forests, though establishment limitation for these species was lowest in the remnant forests. Species were generally seed limited and almost all were establishment limited; these patterns were consistent across treatments. However, our results suggest that differences in recruitment rates could be driven by differential dispersal to treatments with higher canopy cover. We found evidence that barriers to recruitment shift during succession, with the influence of seed limitation, mediated by species-level seed deposition rates, giving way to niche-based processes. However, establishment limitation was lowest in the remnant forests for large-seeded and late successional species, highlighting the importance of habitat specialization and life-history traits in dictating recruitment dynamics. Overall, results demonstrate that active restoration approaches such as tree planting catalyze forest recovery, not only by decreasing components of seed limitation, but also by developing canopy cover that increases establishment rates of larger-seeded species.

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.

© 2015 Society for Ecological Restoration. 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.

Establishing diverse mycorrhizal fungal communities is considered important for forest recovery, yet mycorrhizae may have complex effects on tree growth depending on the composition of fungal species present. In an effort to understand the role of mycorrhizal fungi community in forest restoration in southern Costa Rica, we sampled the arbuscular mycorrhizal fungal (AMF) community across eight sites that were planted with the same species (Inga edulis, Erythrina poeppigiana, Terminalia amazonia, and Vochysia guatemalensis) but varied twofold to fourfold in overall tree growth rates. The AMF community was measured in multiple ways: as percent colonization of host tree roots, by DNA isolation of the fungal species associated with the roots, and through spore density, volume, and identity in both the wet and dry seasons. Consistent with prior tropical restoration research, the majority of fungal species belonged to the genus Glomus and genus Acaulospora, accounting for more than half of the species and relative abundance found on trees roots and over 95% of spore density across all sites. Greater AMF diversity correlated with lower soil organic matter, carbon, and nitrogen concentrations and longer durations of prior pasture use across sites. Contrary to previous literature findings, AMF species diversity and spore densities were inversely related to tree growth, which may have arisen from trees facultatively increasing their associations with AMF in lower soil fertility sites. Changes to AMF community composition also may have led to variation in disturbance susceptibility, host tree nutrient acquisition, and tree growth. These results highlight the potential importance of fungal-tree-soil interactions in forest recovery and suggest that fungal community dynamics could have important implications for tree growth in disturbed soils.

Restoring small-scale habitat heterogeneity in highly diverse systems, like tropical forests, is a conservation challenge and offers an excellent opportunity to test factors affecting community assembly. We investigated whether (1) the applied nucleation restoration strategy (planting tree islands) resulted in higher habitat heterogeneity than more homogeneous forest restoration approaches, (2) increased heterogeneity resulted in more diverse tree recruitment, and (3) the mean or coefficient of variation of habitat variables best explained tree recruitment. We measured soil nutrients, overstory and understory vegetation structure, and tree recruitment at six sites with three 5- to 7-year-old restoration treatments: control (no planting), planted tree islands, and conventional, mixed-species tree plantations. Canopy openness and soil base saturation were more variable in island treatments than in controls and plantations, whereas most soil nutrients had similar coefficients of variation across treatments, and bare ground was more variable in control plots. Seedling and sapling species density were equivalent in plantations and islands, and were substantially higher than in controls. Species spatial turnover, diversity, and richness were similar in island and plantation treatments. Mean canopy openness, rather than heterogeneity, explained the largest proportion of variance in species density. Our results show that, whereas canopy openness and soil base saturation are more heterogeneous with the applied nucleation restoration strategy, this pattern does not translate into greater tree diversity. The lack of a heterogeneity-diversity relationship is likely due to the fact that recruits respond more strongly to mean resource gradients than variability at this early stage in succession, and that seed dispersal limitation likely reduces the available species pool. Results show that planting tree islands facilitates tree recruitment to a similar degree as intensive plantation-style restoration strategies.