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Forest restoration is increasingly heralded as a global strategy to conserve biodiversity and mitigate climate change, yet long-term studies that compare the effects of different restoration strategies on tree recruit demographics are lacking. We measured tree recruit survival and growth annually in three restoration treatments—natural regeneration, applied nucleation and tree plantations—replicated at 13 sites in southern Costa Rica—and evaluated the changes over a decade. Early-successional seedlings had 14% higher survival probability in the applied nucleation than natural regeneration treatments. Early-successional sapling growth rates were initially 227% faster in natural regeneration and 127% faster in applied nucleation than plantation plots but converged across restoration treatments over time. Later-successional seedling and sapling survival were similar across treatments but later-successional sapling growth rates were 39% faster in applied nucleation than in plantation treatments. Results indicate that applied nucleation was equally or more effective in enhancing survival and growth of naturally recruited trees than the more resource-intensive plantation treatment, highlighting its promise as a restoration strategy. Finally, tree recruit dynamics changed quickly over the 10-year period, underscoring the importance of multi-year studies to compare restoration interventions and guide ambitious forest restoration efforts planned for the coming decades. This article is part of the theme issue ‘Understanding forest landscape restoration: reinforcing scientific foundations for the UN Decade on Ecosystem Restoration’. 

Abstract Reversing large-scale habitat degradation and deforestation goes beyond what can be achieved by site-level ecological restoration and a landscape ecology perspective is fundamental. Here we assess the relative importance of tree cover and its configuration on forest-dependent birds and late-successional tree seedlings in restoration sites in southern Costa Rica. The abundance and species richness of birds increased in landscapes with more corridors, higher tree cover, and lower levels of fragmentation, highlighting the importance of riparian corridors for connectivity, and continuous tree cover as suitable habitat. Landscape variables affected abundance and species richness of seedlings similarly, but effects were weaker, possibly because seedlings face establishment limitation in addition to dispersal limitation. Moreover, the scale of landscape effects on seedlings was small, likely because proximal individual trees can significantly influence recruitment in restoration plots. Results underscore the importance of incorporating landscape-level metrics to restoration projects, as knowing the extent, and how the landscape may affect restoration outcomes can help to infer what kind of species will arrive to restoration plots. 

Choosing appropriate forest restoration interventions is challenging. Natural regeneration can rapidly facilitate forest recovery in many situations. However, barriers such as dispersal limitation and competition with non-native species can require assisted restoration approaches to facilitate plant community recovery. We used a study that has directly compared the outcomes of tropical wet forest restoration interventions across 11 replicate sites in southern Costa Rica. Within this framework, we examined the functional recovery trajectories of recruiting tree sapling communities across a gradient of restoration interventions including low (natural regeneration), intermediate (applied nucleation), and high (plantation) initial resource-investment, which we compared to remnant reference forest. We collated leaf and stem functional traits for tree species that comprised the bulk of recruiting saplings, then determined how community-weighted trait means and functional diversity metrics changed over a decade across treatments. Results show that assisted restoration approaches (applied nucleation, plantation) sped the development of more functionally diverse tree communities, more than tripling the functional richness (FRic) of recruiting communities when compared to natural regeneration. However, functional dispersion (i.e., the trait range of dominant species) was equivalent across interventions, and between 28 and 44% lower than remnant forest, indicating that increases in FRic under assisted restoration were driven by species recruiting in low abundances (<10 individuals across treatments). Recruits in assisted restoration treatments also had 10–15% tougher, less-palatable leaves, and leaves were even tougher in reference forest, which could be driven by increasing herbivory pressure along the gradient of interventions. Results show that tracking simple metrics such as species richness can mask a more mechanistic understanding of ecosystem recovery that is elucidated by taking a functional trait-driven approach toward evaluating outcomes. For example, our work identified a paucity of dense-wooded species recruiting across restoration interventions, wood density was 11–13% lower in restoration treatments than reference forests, underscoring such species as prime targets for enrichment planting. Overall, findings suggest that assisted restoration can catalyze the functional recovery of naturally recruiting tree communities in landscapes that are slow to recover naturally and highlight the importance of evaluating how different components of functional diversity shift over time to fully understand restoration outcomes. 

Vegetation processes are fundamentally limited by nutrient and water availability, the uptake of which is mediated by plant roots in terrestrial ecosystems. While tropical forests play a central role in global water, carbon, and nutrient cycling, we know very little about tradeoffs and synergies in root traits that respond to resource scarcity. Tropical trees face a unique set of resource limitations, with rock-derived nutrients and moisture seasonality governing many ecosystem functions, and nutrient versus water availability often separated spatially and temporally. Root traits that characterize biomass, depth distributions, production and phenology, morphology, physiology, chemistry, and symbiotic relationships can be predictive of plants’ capacities to access and acquire nutrients and water, with links to aboveground processes like transpiration, wood productivity, and leaf phenology. In this review, we identify an emerging trend in the literature that tropical fine root biomass and production in surface soils are greatest in infertile or sufficiently moist soils. We also identify interesting paradoxes in tropical forest root responses to changing resources that merit further exploration. For example, specific root length, which typically increases under resource scarcity to expand the volume of soil explored, instead can increase with greater base cation availability, both across natural tropical forest gradients and in fertilization experiments. Also, nutrient additions, rather than reducing mycorrhizal colonization of fine roots as might be expected, increased colonization rates under scenarios of water scarcity in some forests. Efforts to include fine root traits and functions in vegetation models have grown more sophisticated over time, yet there is a disconnect between the emphasis in models characterizing nutrient and water uptake rates and carbon costs versus the emphasis in field experiments on measuring root biomass, production, and morphology in response to changes in resource availability. Closer integration of field and modeling efforts could connect mechanistic investigation of fine-root dynamics to ecosystem-scale understanding of nutrient and water cycling, allowing us to better predict tropical forest-climate feedbacks. 

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.

 

In degraded tropical landscapes, lack of seed dispersal can strongly limit recovery, and restoration interventions can overcome this barrier by attracting dispersers. However, seed dispersal patterns are typically studied over short time periods, thus the influences of temporal and spatial variability on seed arrival cannot be teased apart.

The choice of management approach can have important implications for restoration‐mediated community reassembly. Accordingly, we used a 3.5‐year record of seed deposition in pre‐montane tropical wet forest in southern Costa Rica to examine how seed arrival differed between passive (natural regeneration) and active (applied nucleation, plantation) restoration after a decade of recovery, compared to remnant forest. We investigated: (a) how restoration treatments affected seed deposition rates and community composition; (b) how within‐plot heterogeneity of animal‐dispersed seed deposition varied by intervention; and (c) how interannual variation influenced animal‐dispersed seed arrival across treatments.

Overall seed rain composition and diversity in restoration treatments was converging towards, but still differed substantially from, remnant forest (89.7%, 86.6% and 76.3% Shannon diversity recovered in applied nucleation, plantation and natural regeneration respectively).

Within‐plot animal‐dispersed seed heterogeneity was similar in applied nucleation and remnant forest, 27.0% more heterogeneous in applied nucleation than plantation, and equivalent when comparing natural regeneration to either applied nucleation or plantation.

In contrast to active interventions, animal‐dispersed tree and shrub communities did not differ year to year in natural regeneration, which may promote the assembly of relatively homogeneous plant communities at this successional stage.

Synthesis and applications. Compared to natural regeneration, active restoration interventions: (a) catalysed the recovery of seed diversity (overall Shannon diversity 17.5% and 13.4% higher in applied nucleation and plantation respectively), (b) shifted seed community composition towards remnant forest more rapidly (overall Shannon diversity 13.4% and 10.2% closer), (c) almost doubled the proportion of later‐successional tree species arriving, and (d) had seed communities that differed year to year—a pattern not observed in natural regeneration. Finally, applied nucleation was the only intervention where seed arrival was as spatially heterogeneous as remnant forest, highlighting that this approach may facilitate the recovery of specific natural dispersal processes.