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Abstract The effects of vertebrate seed predation on the regeneration of restored forests are not well understood because most past studies have focused on seed predation within the first few years after restoration and have measured seed removal without quantifying subsequent seedling establishment of seeds that avoid predation. Quantifying the establishment of seeds that escape predation in restored forests at later stages of regrowth is crucial for anticipating longer‐term recovery trajectories. Here, we evaluated the potential role of vertebrate seed predators in limiting recruitment of later‐successional tree species in nine forests actively restored ≥15 years prior and in four paired remnant forest fragments embedded in an agricultural landscape in southern Costa Rica. We conducted seed addition experiments with four tree species inside and outside vertebrate exclosures and used camera trapping to detect seed predators. To determine the fate of seeds that avoided predation, we also measured seedling establishment after 1 year, given that other mortality factors may compensate in the absence of vertebrate seed predation. We detected two species of birds and five species of granivorous mammals removing seeds. Seed tagging indicated that most removal resulted in predation. For three of the four tree species tested, vertebrate seed predation reduced seedling establishment. The magnitude of this effect depended on species' susceptibility to other causes of mortality during the seed‐to‐seedling transition. Our study demonstrates that vertebrate seed predators can substantially reduce later‐successional seedling recruitment in restored forests and should be considered alongside dispersal limitation and microsite conditions as factors slowing forest recovery. Abstract in Spanish is available with online material. 

Abstract Tropical forest restoration presents a potential lifeline to mitigate climate change and biodiversity crises in the Anthropocene. Yet, the extent to which human interventions, such as tree planting, accelerate the recovery of mature functioning ecosystems or redirect successional trajectories toward novel states remains uncertain due to a lack of long‐term experiments. In 2004–2006, we established three 0.25‐ha plots at 10 sites in southern Costa Rica to test three forest restoration approaches: natural regeneration (no planting), applied nucleation (planting in patches), and plantation (full planting). In a comprehensive survey after 16–18 years of recovery, we censused >80,000 seedlings, saplings, and trees from at least 255 species across 26 restoration plots (nine natural regeneration, nine applied nucleation, eight plantation) and six adjacent reference forests to evaluate treatment effects on recruitment patterns and community composition. Both applied nucleation and plantation treatments resulted in significantly elevated seedling and sapling establishment and more predictable community composition compared with natural regeneration. Similarity of vegetation composition to reference forest tended to scale positively with treatment planting intensity. Later‐successional species with seeds ≥5 mm had significantly greater seedling and sapling abundance in the two planted treatments, and plantation showed similar recruitment densities of large‐seeded (≥10 mm) species to reference forest. Plantation tended toward a lower abundance of early‐successional recruits than applied nucleation. Trees (≥5 cm dbh) in all restoration treatments continued to be dominated by a few early‐successional species and originally transplanted individuals. Seedling recruits of planted taxa were more abundant in applied nucleation than the other treatments though few transitioned into the sapling layer. Overall, our findings show that active tree planting accelerates the establishment of later‐successional trees compared with natural regeneration after nearly two decades. While the apparent advantages of higher density tree planting on dispersal and understory establishment of larger seeded, later‐successional species recruitment is notable, more time is needed to assess whether these differences will persist and transition to the more rapid development of a mature later‐successional canopy. Our results underscore the need for ecological restoration planning and monitoring that targets biodiversity recovery over multiple decades. 

Applied nucleation and other spatially patterned restoration methods are promising approaches for scaling up projects to meet ambitious international restoration commitments in an ecologically and economically sound manner. Much of the corresponding literature to date, however, has centered around theoretical discussions and small‐scale studies that are largely divorced from constraints faced by restoration practitioners. We briefly review recent academic literature about applied nucleation and other spatially patterned restoration methods and discuss practical challenges to their implementation. We offer several recommendations to move spatially patterned restoration from an academic conversation to scalable application, including: (1) comparing different planting designs and natural regeneration within the same system at an appropriate scale; (2) monitoring ecological outcomes throughout the restored area over sufficient time to evaluate recovery; (3) quantifying costs and documenting other logistical constraints to implementation; and (4) exploring methods for using unplanted areas to provide benefits to landholders until planted vegetation establishes. 

Choosing effective methods to restore habitat for the diverse faunal assemblages of tropical forests is hampered by lack of long-term data comparing multiple restoration treatments. We conducted area counts of bird assemblages over 12 years (~5–17 years since restoration) in a blocked experiment with two active planted treatments (tree plantations and applied nucleation) and a passive restoration treatment (natural regeneration) replicated at 11 sites in Costa Rica. We also surveyed six pastures and five remnant forest sites to assess recovery of avian species richness, composition, forest specialists, and range-restricted species in restoration plots relative to degraded and reference systems. Restoration treatments showed increased resemblance of avian assemblages to remnant forest over time. Applied nucleation proved equally effective as plantation, despite a reduced planted area, whereas natural regeneration recovered more slowly. Assemblage-level trends in avian species richness and compositional similarity to reference forest are underpinned by reductions in use by pasture birds and by gradual increases in richness of forest-affiliated species. Because forest-affiliated species tend to have narrower distributions than the open-country species they replace, forest restoration can reduce biotic homogenization at the local scale. Restoration practitioners should consider applied nucleation as an alternative to standard plantations if seeking rapid recovery of bird assemblages. However, the ecological return on investment from natural regeneration increases over a couple of decades. Managers should monitor trends in forest-affiliated and rangerestricted species to track the recovery of the full avian assemblages, since coarse metrics like species richness and overall compositional similarity may plateau relatively quickly 

With increased interest in forest restoration comes an urgent need to provide accurate, scalable, and cost‐effective monitoring tools. The ubiquity of smartphones has led to a surge in monitoring apps. We reviewed and assessed monitoring apps found through web searches and conversations with practitioners. We identified 42 apps that (1) automatically monitor indicators or (2) facilitate data entry. We selected the five most promising from the first category, based on their relevance, availability, stability, and user support. We compared them to traditional field techniques in a well‐studied restoration project in Costa Rica. We received further feedback from 15 collaborator organizations that evaluated these in their corresponding field restoration sites. Diameter measurements correlated well with traditional tape‐based measurements (R² = 0.86–0.89). Canopy openness and ground cover showed weaker correlations to densiometer and quadrat cover measurements (R² = 0.42–0.51). Apps did not improve labor efficiency but do preclude the purchase of specialized field equipment. The apps reviewed here need further development and validation to support monitoring adequately, especially in the tropics. Estimates of development and maintenance costs, as well as statistics on user uptake, are required for cost‐effective development. We recommend a coordinated effort to develop dedicated restoration monitoring apps that can speed up and standardize the collection of indicators and provide evidence on restoration outcomes alongside a centralized repository of this information. 

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’. 

Insect herbivory is one of the major drivers of seedling mortality in the tropics and influences plant abundances and community composition. Anthropogenic disturbance can alter patterns of insect herbivory with potential consequences on plant communities in restored forests. We planted seedlings of early‐ and later‐stage successional tree species in 13–15‐year‐old restored and remnant tropical forests. We then either excluded insect herbivores or left seedlings exposed to examine how insect herbivory‐affected seedling mortality. Early‐successional seedlings experienced similar decreases in mortality when insect herbivores were excluded from both restored and remnant forest sites, but this effect was smaller and driven by only a few species in restored forests. Later‐successional seedlings experienced a stronger decrease in mortality between open and insect‐excluded treatments in remnant than restored sites. Our results suggest that herbivory‐driven seedling mortality is lower in restored forests, particularly for later‐successional seedlings. Results are encouraging from a restoration perspective because recruitment of later‐successional seedlings is a key component of ecosystem recovery. However, if reductions in seedling mortality continue over the long term, this may affect tree community composition as succession progresses.