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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.