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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. [ABSTRACT FROM AUTHOR]

Questions Vascular epiphytes constitute a large proportion of tropical forest plant biodiversity, but are among the slowest plants to recolonize secondary forests. We asked whether tree planting for ecological restoration accelerates epiphyte community recovery. Does the spatial configuration of tree planting matter? What landscape contexts are most suitable for epiphyte restoration? Location Restored pastures in premontane Coto Brus County, Puntarenas, Costa Rica. Methods We surveyed vascular epiphyte species growing on the lower trunks of 1083 trees in 13 experimental restoration sites. Each site contained three 0.25-ha treatment plots: natural regeneration, trees planted in patches or 'islands' and tree plantations. Sites spanned elevational (1100-1430 m) and deforestation (4-94% forest cover within a 100-m radius around each site) gradients. Results Vascular epiphytes were twice as diverse in planted restoration plots (islands and plantations) as in natural regeneration; we observed this at the scale of individual host trees and within 0.25-ha treatment plots. Contributing factors included that trees in planted restoration plots were larger, older, more abundant and composed of different species than trees in naturally regenerating plots. Epiphyte species richness increased with surrounding forest cover within 100-150 m of restoration plots. Epiphyte communities were also twice as diverse at higher (1330-1430 m) vs lower (1100-1290 m) elevation sites. Epiphyte groups responded differently to restoration treatments and landscape factors; ferns were responsible for higher species richness in planted restoration plots, whereas angiosperms drove elevation and forest cover effects. Conclusions Tree planting for ecological restoration enriched epiphyte communities compared to natural regeneration, likely because planted forests contained more, bigger and older trees. Tree island plantings were equally effective compared to larger and more expensive plantations. Restoration sites nearer to existing forests had richer epiphyte recolonization, likely because nearby forests provisioned restoration sites with angiosperm seeds. Collectively, results suggest that restoration practitioners can enrich epiphyte community development by planting trees in areas with higher surrounding forest cover, particularly at higher elevations. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Understanding tropical biology is important for solving complex problems such as climate change, biodiversity loss, and zoonotic pandemics, but biology curricula view research mostly via a temperate-zone lens. Integrating tropical research into biology education is urgently needed to tackle these issues. Tropical biology is currently largely absent from undergraduate curricula. The integration of authentic research in tropical biology could promote internationality among scientists and provide experiential learning that enables students to address complex global problems. [ABSTRACT FROM AUTHOR]

Abstract: We investigated the hypothesis that maritime climatic factors associated with summer fog and low cloud stratus (summer marine layer) help explain the compositional diversity of chaparral in the coast range of central California. We randomly sampled chaparral species composition in 0.1‐hectare plots along a coast‐to‐interior gradient. For each plot, climatic variables were estimated and soil samples were analyzed. We used Cluster Analysis and Principle Components Analysis to objectively categorize plots into climate zone groups. Climate variables, vegetation composition and various diversity measures were compared across climate zone groups using ANOVA and nonmetric multidimensional scaling. Differences in climatic variables that relate to summer moisture availability and winter freeze events explained the majority of variance in measured conditions and coincided with three chaparral assemblages: maritime (lowland coast where the summer marine layer was strongest), transition (upland coast with mild summer marine layer influence and greater winter precipitation), and interior sites that generally lacked late summer water availability from either source. Species turnover (β‐diversity) was higher among maritime and transition sites than interior sites. Coastal chaparral differs from interior chaparral in having a higher obligate seeder to facultative seeder (resprouter) ratio and by being dominated by various Arctostaphylos species as opposed to the interior dominant, Adenostoma fasciculatum. The maritime climate influence along the California central coast is associated with patterns of woody plant composition and β‐diversity among sites. Summer fog in coastal lowlands and higher winter precipitation in coastal uplands combine to lower late dry season water deficit in coastal chaparral and contribute to longer fire return intervals that are associated with obligate seeders and more local endemism. Soil nutrients are comparatively less important in explaining plant community composition, but heterogeneous azonal soils contribute to local endemism and promote isolated chaparral patches within the dominant forest vegetation along the coast. [ABSTRACT FROM AUTHOR]

1. Active restoration to rehabilitate degraded tropical lands often involves planting tree seedlings, an effective but expensive approach if large areas are planted. Planting small patches of vegetation (tens to a few hundred square metres) has recently been suggested as a more economical restoration technique that mimics natural regeneration processes. However, few studies have examined the consequences of restoration patch size on animals, whose presence and activities are often key to successful ecosystem recovery. 2. We examined the effects of patch size on the foraging behaviour of four resident tropical bird species in a replicated forest restoration experiment in southern Costa Rica. We also measured arthropod abundance and anti-predator vigilance behaviour to assess whether variation in food availability or predation risk could explain patch size effects on foraging behaviour. 3. Prey attack rates were highest, and the effort required to find prey was lowest, in larger patches for three of the four bird species. Arthropod density was approximately twice as great in larger patches (>3500 m2) compared with smaller patches (<350 m2). Evidence for patch size differences in predation risk was more limited but risk may be higher in smaller patches. The results indicate that food availability is the primary mechanism driving patch size effects on foraging behaviour, with predation risk being an additional influence for some species in some years. 4. Synthesis and applications. As demonstrated in this study, patches of tens to a few hundreds of metres squared are likely to provide fewer food resources and potentially less cover from predators for vertebrates that use woody habitat, compared with patches of a few thousand square metres. The more limited resources in smaller patches are likely to have short-term and, potentially, long-term consequences for the fitness of organisms. When considering restoration project design, the potential economic and other benefits of planting in smaller patches must be weighed with the potentially negative ecological effects on some taxonomic groups. To increase the probability that patches provide adequate habitat for the largest number of species, we recommend that when financial resources are available, patches of at least a few thousand square metres be planted. [ABSTRACT FROM AUTHOR]

Remnant trees and forest fragments in agricultural landscapes can be important sources of propagules to facilitate forest recovery. However, many studies simply quantify forest cover in the surrounding landscape as a percentage, with little attention given to species composition, and subsequently fail to detect an effect on recruitment patterns. We assessed the relative importance of the spatial distribution and life‐history traits of 77 tree species on recruitment patterns at a landscape scale in a well‐replicated long‐term restoration study in southern Costa Rica. We censused and mapped potential mother trees in a 100‐m buffer surrounding eight replicate restoration plots and quantified respective tree recruits within each plot. We assessed how mother tree abundance, species life‐history characteristics (seed size, dispersal mode), tree size (DBH, height) and distance to restoration plot affected recruitment at coarse (plot: 50 × 50 m) and fine (quadrat: 3 × 3 m) spatial scales. The presence of a mother tree within 100 m of a restoration plot resulted in a 10‐fold increase in potential mean recruitment. Mother tree abundance was also an important driver of recruit density, and particularly so for large‐seeded (≥ 5 mm) zoochorous species with a fivefold increase in recruit density across the observed mother tree abundance range. An interaction between mother tree abundance and proximity demonstrated that the effect of mother tree abundance on recruit density was important but waned with increasing distance from restoration plots. At the fine spatial scale, proximity was uniformly important; height and DBH of the closest potential mother tree also affected recruit abundance but responses differed by seed size. Results highlight the importance of remnant vegetation composition to the recovery of adjacent degraded habitats, underscoring the outsized role nearby remnant forest and isolated trees can play for the persistence of localized biodiversity. [ABSTRACT FROM AUTHOR]

Given that few ecosystems on the Earth have been unaffected by humans, restoring them holds great promise for stemming the biodiversity crisis and ensuring ecosystem services are provided to humanity. Nonetheless, few studies have documented the recovery of ecosystems globally or the rates at which ecosystems recover. Even fewer have addressed the added benefit of actively restoring ecosystems versus allowing them to recover without human intervention following the cessation of a disturbance. Our metaanalysis of 400 studies worldwide that document recovery from large-scale disturbances, such as oil spills, agriculture and logging, suggests that though ecosystems are progressing towards recovery following disturbances, they rarely recover completely. This result reinforces conservation of intact ecosystems as a key strategy for protecting biodiversity. Recovery rates slowed down with time since the disturbance ended, suggesting that the final stages of recovery are the most challenging to achieve. Active restoration did not result in faster or more complete recovery than simply ending the disturbances ecosystems face. Our results on the added benefit of restoration must be interpreted cautiously, because few studies directly compared different restoration actions in the same location after the same disturbance. The lack of consistent value added of active restoration following disturbance suggests that passive recovery should be considered as a first option; if recovery is slow, then active restoration actions should be better tailored to overcome specific obstacles to recovery and achieve restoration goals.We call for a more strategic investment of limited restoration resources into innovative collaborative efforts between scientists, local communities and practitioners to develop restoration techniques that are ecologically, economically and socially viable. [ABSTRACT FROM AUTHOR]

Dr. Emilio M. Bruno is not included in the author byline. He should be listed as the fourth author, and his affiliation is: Center for Latin American Studies, University of Florida, Gainesville, Florida, and Department of Wildlife Ecology & Conservation, University of Florida, Gainesville, Florida, United States of America. [Extracted from the article]

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. [ABSTRACT FROM AUTHOR]

Birds both promote and prosper from forest restoration. The ecosystem functions birds perform can increase the pace of forest regeneration and, correspondingly, increase the available habitat for birds and other forest-dependent species. The aim of this study was to learn how tropical forest restoration treatments interact with landscape tree cover to affect the structure and composition of a diverse bird assemblage. We sampled bird communities over two years in 13 restoration sites and two old-growth forests in southern Costa Rica. Restoration sites were established on degraded farmlands in a variety of landscape contexts, and each included a 0.25-ha plantation, island treatment (trees planted in patches), and unplanted control. We analyzed four attributes of bird communities including frugivore abundance, nectarivore abundance, migrant insectivore richness, and compositional similarity of bird communities in restoration plots to bird communities in old-growth forests. All four bird community variables were greater in plantations and/or islands than in control treatments. Frugivore and nectarivore abundance decreased with increasing tree cover in the landscape surrounding restoration plots, whereas compositional similarity to old-growth forests was greatest in plantations embedded in landscapes with high tree cover. Migrant insectivore richness was unaffected by landscape tree cover. Our results agree with previous studies showing that increasing levels of investment in active restoration are positively related to bird richness and abundance, but differences in the effects of landscape tree cover on foraging guilds and community composition suggest that trade-offs between biodiversity conservation and bird-mediated ecosystem functioning may be important for prioritizing restoration sites. [ABSTRACT FROM AUTHOR]

Many terrestrial plant communities, especially forests, have been shown to lag in response to rapid climate change. Grassland communities may respond more quickly to novel climates, as they consist mostly of short-lived species, which are directly exposed to macroclimate change. Here we report the rapid response of grassland communities to climate change in the California Floristic Province. We estimated 349 vascular plant species' climatic niches from 829,337 occurrence records, compiled 15 long-term community composition datasets from 12 observational studies and 3 global change experiments, and analysed community compositional shifts in the climate niche space. We show that communities experienced significant shifts towards species associated with warmer and drier locations at rates of 0.0216 ± 0.00592 °C yr - 1 (mean ± s.e.) and -3.04 ± 0.742 mm yr - 1 , and these changes occurred at a pace similar to that of climate warming and drying. These directional shifts were consistent across observations and experiments. Our findings contrast with the lagged responses observed in communities dominated by long-lived plants and suggest greater biodiversity changes than expected in the near future., (© 2024. The Author(s).)

Livestock grazing is often used as a land management tool to maximize vegetation diversity in grassland ecosystems worldwide. Prior research has shown that cattle grazing benefits native annual forb species in California's coastal prairies, but drought and increasing aridity may alter this relationship. In 2016 and 2017, we resurveyed the vegetation structure, native annual forb cover, and native annual forb richness in ten grazed and ungrazed prairies that were originally measured in 2000 and 2001 along a 200-km gradient from Monterey to Sonoma counties in California. We found that grazed prairies continued to have significantly lower vegetation height and thatch depth than ungrazed prairies, and that shrub encroachment over the 15-year period was significantly greater in ungrazed prairies. Furthermore, grazed prairies continued to have greater native annual forb richness (4.9 species per site) than ungrazed sites (3.0 species per site), but native annual forb richness declined by 2.8 species per site in grazed prairies and 0.1 species per site in ungrazed prairies between survey periods. We suggest that severe drought and increasing aridity may be driving declines in native annual forb richness in grazed prairies. The species we recorded only in earlier surveys were disproportionately wetland-associated and had higher average specific leaf area than species that remained through the second survey period. Finally, the cover of native annual species increased regardless of whether prairies were grazed, suggesting that the high precipitation in 2017 may have benefitted the native annual forb species that persisted at sites between surveys. Our study shows that weather conditions affect the outcomes of land management strategies., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2022 Lesage et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Extensive evidence shows that regional (gamma) diversity is often lower across restored landscapes than in reference landscapes, in part due to common restoration practices that favor widespread species through selection of easily-grown species with high survival and propagation practices that reduce genetic diversity. We discuss approaches to counteract biotic homogenization, such as reintroducing species that are adapted to localized habitat conditions and are unlikely to colonize naturally; periodically reintroducing propagules from remnant populations to increase genetic diversity; and reintroducing higher trophic level fauna to restore interaction networks and processes that promote habitat heterogeneity. Several policy changes would also increase regional diversity; these include regional coordination amongst restoration groups, financial incentives to organizations producing conservation-valued species, and experimental designations for rare species introductions., Competing Interests: Declarations of interest No interests are declared., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Competing Interests: The authors have declared that no competing interests exist.

To constrain global warming, we must strongly curtail greenhouse gas emissions and capture excess atmospheric carbon dioxide 1,2 . Regrowing natural forests is a prominent strategy for capturing additional carbon 3 , but accurate assessments of its potential are limited by uncertainty and variability in carbon accumulation rates 2,3 . To assess why and where rates differ, here we compile 13,112 georeferenced measurements of carbon accumulation. Climatic factors explain variation in rates better than land-use history, so we combine the field measurements with 66 environmental covariate layers to create a global, one-kilometre-resolution map of potential aboveground carbon accumulation rates for the first 30 years of natural forest regrowth. This map shows over 100-fold variation in rates across the globe, and indicates that default rates from the Intergovernmental Panel on Climate Change (IPCC) 4,5 may underestimate aboveground carbon accumulation rates by 32 per cent on average and do not capture eight-fold variation within ecozones. Conversely, we conclude that maximum climate mitigation potential from natural forest regrowth is 11 per cent lower than previously reported 3 owing to the use of overly high rates for the location of potential new forest. Although our data compilation includes more studies and sites than previous efforts, our results depend on data availability, which is concentrated in ten countries, and data quality, which varies across studies. However, the plots cover most of the environmental conditions across the areas for which we predicted carbon accumulation rates (except for northern Africa and northeast Asia). We therefore provide a robust and globally consistent tool for assessing natural forest regrowth as a climate mitigation strategy.

Global forest restoration targets have been set, yet policy makers and land managers lack guiding principles on how to invest limited resources to achieve them. We conducted a meta-analysis of 166 studies in naturally regenerating and actively restored forests worldwide to answer: (1) To what extent do floral and faunal abundance and diversity and biogeochemical functions recover? (2) Does recovery vary as a function of past land use, time since restoration, forest region, or precipitation? (3) Does active restoration result in more complete or faster recovery than passive restoration? Overall, forests showed a high level of recovery, but the time to recovery depended on the metric type measured, past land use, and region. Abundance recovered quickly and completely, whereas diversity recovered slower in tropical than in temperate forests. Biogeochemical functions recovered more slowly after agriculture than after logging or mining. Formerly logged sites were mostly passively restored and generally recovered quickly. Mined sites were nearly always actively restored using a combination of planting and either soil amendments or recontouring topography, which resulted in rapid recovery of the metrics evaluated. Actively restoring former agricultural land, primarily by planting trees, did not result in consistently faster or more complete recovery than passively restored sites. Our results suggest that simply ending the land use is sufficient for forests to recover in many cases, but more studies are needed that directly compare the value added of active versus passive restoration strategies in the same system. Investments in active restoration should be evaluated relative to the past land use, the natural resilience of the system, and the specific objectives of each project., Competing Interests: The authors have declared that no competing interests exist.

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.

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.

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.

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.

Tropical forest restoration strategies have the potential to accelerate the recovery of the nutrient cycles in degraded lands. Litter production and its decomposition represent the main transfer of organic material and nutrients into the soil substrate. We evaluated litter production, accumulation on the forest floor, and its decomposition under three restoration strategies: plantation (entire area planted with trees), island (trees planted in patches of three different sizes) and control (natural regeneration) plots. We also compared restoration strategies to young secondary forest (7-9 yr). Restoration treatments were established in 50 x 50m plots in June 2004 at six sites in Southern Costa Rica. Planted tree species included two native timber species (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N fixers (Erythrina poeppigiana and Inga edulis). Litter was collected every 15 days between September 2008 and August 2009 in 12 0.25m2 litter traps distributed within each plot; litter that accumulated on the soil surface was collected at four locations (0.25m2 quadrats) within each plot in February and May 2009. Total litter production in plantation (6.3Mg/ha) and secondary forest (7.3Mg/ha) did not differ, but were greater than in islands (3.5Mg/ha) and control (1.4 Mg/ha). Plantation had greatest accumulation of litter on the soil surface (10.6 Mg/ha) as compared to the other treatments (SF = 7.2; I = 6.7; C = 4.9). Secondary forest was the only treatment with a greater annual production of litter than litter accumulation on the soil surface. Carbon storage in litter was similar between plantation and secondary forest, and significantly greater than the other treatments. No differences were found for carbon concentration and storage in the soil among treatments. There was also high variability in the production and accumulation of litter and carbon among sites. Active restoration treatments accelerated the production of litter and carbon storage in comparison to areas under natural recovery. However, the nutrient cycle has not necessarily been restored under these conditions, as high litter accumulation on the soil surface indicates a low decomposition rate, which slows nutrient return to the soil.

Introducing rare plants to new sites for conservation to offset effects of habitat destruction requires detailed knowledge of habitat requirements, plant demography, and management needs. We conducted a factorial experiment replicated at three coastal prairie sites to test the effects of clipping frequency and litter accumulation on seed germination, seedling survival, reproduction, and seedling recruitment of introduced populations of the endangered, tall-stature, annual forb, Holocarpha macradenia (DC.) E. Greene. Clipping favored H. macradenia, primarily by enhancing seed germination and flower production. Litter accumulation had no effect on seed germination, even after 5 years of treatments. Seedling recruitment was highly site specific with large numbers of recruits recorded at only one of three sites. Although recruitment of seedlings was higher in clipped plots for 2-3 years, by 4-5 years after introduction very few seedlings survived to reproduction in any treatment. We attribute this result to a combination of poor habitat quality, small population size, and lack of a seed bank. We were unsuccessful in introducing this relatively well-studied species of concern to apparently suitable habitat at multiple sites in multiple years, which suggests that translocating rare plant populations to mitigate for habitat destruction is an expensive and highly uncertain endeavor.

Only recently have studies addressed the effect of early-colonizing vegetation on tree seedling survival and growth during secondary succession in tropical old fields, and few studies have elucidated the physiological responses of tree seedlings to different vegetational communities. We compared growth and various photosynthetic parameters for seedlings of four rain-forest tree species, Cedrela tonduzii, Inga punctata, Ocotea whitei, and Tapirira mexicana, growing in areas of pasture grass and shrubs in early-successional abandoned pasture in Costa Rica; in addition, we made measurements for two species in forest gaps. We tested the general hypothesis that early-colonizing shrubs facilitate growth of forest tree seedlings. Specifically, we measured microclimate, growth, CO 2 assimilation, stomatal conductance, photosystem II quantum yield (ΦPSII), and xanthophyll pigment pools for all seedlings. Photosynthetic flux density (PFD) was higher under grass than shrubs or forest gaps, but was highly variable in each growth environment. For three of the four species, height growth was greatest in the grass compared to the shrubs and forest gaps; growth was similar below grass and shrubs for O. whitei. Photosynthetic capacity, apparent quantum yield, and stomatal conductance did not vary across habitats, but light compensation point and PFD at light saturation tended to be higher in the grass compared to forest and shrub growth environments. Water use efficiency differed across growth environments for three of the species. For plants in ambient PFD and dark-adapted plants, the efficiency of excitation energy transfer through PSII was lowest for plants in the grass compared to shrubs and forest gaps and also differed across species. Measurement of steady-state responses of ΦPSII to increasing PFD indicated a significant effect of growth environment at low PFD for all species and significant effects at high PFD only for I. punctata. All species exhibited a high degree of midday xanthophyll de-epoxidation in the different growth environments. Xanthophyll pigment pool size on an area basis was highest in the grass compared to shrubs and forest gaps for all four species. The results suggest that shrubs do not provide a facilitative effect for growth or photosynthesis for ~1.5-year-old seedlings of these four species. We conclude that site differences in success of tree seedlings during succession are a result of complex interactions of facilitation and competition and are not simply based on physiological responses to PFD.