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QUESTIONS: Natural regeneration is increasingly recognized as a potentially cost‐effective strategy to reach ambitious forest landscape restoration targets, but rates of recovery are notoriously variable. We asked how well initial habitat conditions after cessation of agriculture predict forest recovery after nearly a decade. We aimed to provide land managers with general rules of thumb to assess when it is necessary to invest resources in active restoration, such as tree planting, to accelerate forest recovery. LOCATION: Coto Brus County, Puntarenas, Costa Rica. METHODS: We compiled data on initial vegetation structure, soil nutrients, prior land‐use history and surrounding forest cover at 13 sites. After 8.5 years, we measured vegetation indicators commonly used to assess forest recovery, namely amount of canopy closure and number and diversity of woody recruits. RESULTS: Two variables, grass cover and canopy closure, measured 1.5 years after site abandonment, explained 47–87% of five of the six response variables after 8.5 years; recovery was faster in sites with lower grass cover and higher canopy closure initially. Waiting an additional year to measure initial vegetation variables did not improve model fit. Time since the original forest was cleared explained 62% of change in canopy cover, whereas percentage of surrounding forest cover, length of pasture use and soil variables explained minimal additional variation. CONCLUSIONS: Our results suggest that two easily measurable vegetation variables can provide guidance to land managers and policy makers about where to invest scarce restoration resources to facilitate forest recovery.

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 meta-analysis 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.

How can ambitious forest restoration targets be implemented on the ground?

Monitoring protocols are needed to evaluate the millions of hectares of land that are being proposed for forest restoration in the coming decades. Standardized proposals are critical to evaluate efficacy of restoration strategies, identify triggers for corrective actions, compare results across projects, and generally learn from past projects to inform future restoration efforts. We describe an iterative process, including over 200 stakeholders, to develop a protocol for monitoring Brazilian Atlantic Forest restoration. We give an overview of the ecological, socioeconomic, and management criteria, indicators, and metrics included in the protocol. Strengths of the protocol include the following: (a) testing and use across sites with a range of ages, forest types, past land uses, restoration techniques, and implementing institutions; (b) participation by a broad range of government, nongovernment, private, and academic institutions in the protocol development process; and (c) inclusion of socioeconomic and management criteria. Next steps for facilitating the broad adoption of the protocol in the Atlantic Forest region include providing in person and online training courses, establishing an online repository for storing and comparing monitoring data, and developing smartphone applications to facilitate data collection. Although the protocol was developed for the Atlantic Forest context and further refinements are needed, we think that the Atlantic Forest Pact monitoring protocol may serve as a model to inform the development of similar protocols in other regions, which ultimately could be integrated to produce a pantropical protocol for common use by several restoration forest programs worldwide.

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.

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.

The selection of ecological indicators is an important step toward more effective restoration monitoring. The debate between Reid (2015) and Durigan and Suganuma (2015) regarding the usefulness of species composition for monitoring restoration trajectory is timely and salient, but it lacks a middle way proposal to balance ecological relevance and practical viability. We propose a way forward to resolving this debate, namely using easily measurable functional traits, a type of compositional measure, as an indicator. Assessing functional composition trajectory may help overcome some limitations with taxonomic identification and provide more meaningful outcomes to evaluate restoration success.

© 2015 Springer Science+Business Media Dordrecht Planted woodland islets act as sources of seed that may accelerate woodland development in extensive agricultural landscapes. We assessed a 1-ha plot that was planted with 16 100-m2 islets of holm oak Quercus ilex subsp. ballota seedlings near Toledo (Spain) in 1993. In spring 2014 we measured (1) acorn predation and (2) seedling emergence from seeded acorns at different distances from and orientations around the islets with half of the acorns protected to prevent predation, (3) survival of emerged seedlings, and (4) natural tree establishment outside of the planted islets. Most (96.9 %) unprotected acorns were removed or predated. Seedling emergence from protected acorns ranged from 42.9 % on the northern side of the islets to 13.2 % on the southern side, suggesting a less stressful microclimate on the northern side. Survival of naturally established seedlings was 28.6 % by the end of first summer; seedling mortality was chiefly due to drought (45.0 %) and herbivory (35.0 %). Density of emerged seedlings, surviving seedlings after first dry season, and established oaks >1-year old was similar at different distances from the islets. Over the 21 year time period, 58 new oak individuals >1-year old have established (an average of 3.3 established individuals per ha per year) at an average distance of 6.3 ± 5.4 m away from the closest islet. We conclude that initial oak regeneration triggered by small planted islets in Mediterranean abandoned farmland is slowed down by high acorn predation, seedling herbivory, and stressful microclimatic conditions. Regardless, these islets are a viable tool for regeneration of Mediterranean oak woodland.

© 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.

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.

Plant functional traits, in particular specific leaf area (SLA), wood density and seed mass, are often good predictors of individual tree growth rates within communities. Individuals and species with high SLA, low wood density and small seeds tend to have faster growth rates. If community-level relationships between traits and growth have general predictive value, then similar relationships should also be observed in analyses that integrate across taxa, biogeographic regions and environments. Such global consistency would imply that traits could serve as valuable proxies for the complex suite of factors that determine growth rate, and, therefore, could underpin a new generation of robust dynamic vegetation models. Alternatively, growth rates may depend more strongly on the local environment or growth-trait relationships may vary along environmental gradients. We tested these alternative hypotheses using data on 27 352 juvenile trees, representing 278 species from 27 sites on all forested continents, and extensive functional trait data, 38% of which were obtained at the same sites at which growth was assessed. Data on potential evapotranspiration (PET), which summarizes the joint ecological effects of temperature and precipitation, were obtained from a global data base. We estimated size-standardized relative height growth rates (SGR) for all species, then related them to functional traits and PET using mixed-effect models for the fastest growing species and for all species together. Both the mean and 95th percentile SGR were more strongly associated with functional traits than with PET. PET was unrelated to SGR at the global scale. SGR increased with increasing SLA and decreased with increasing wood density and seed mass, but these traits explained only 3.1% of the variation in SGR. SGR-trait relationships were consistently weak across families and biogeographic zones, and over a range of tree statures. Thus, the most widely studied functional traits in plant ecology were poor predictors of tree growth over large scales. Synthesis. We conclude that these functional traits alone may be unsuitable for predicting growth of trees over broad scales. Determining the functional traits that predict vital rates under specific environmental conditions may generate more insight than a monolithic global relationship can offer. The most widely studied functional traits in plant ecology, specific leaf area, wood density and seed mass, were only weakly associated with tree growth rates over broad scales. Assessing trait-growth relationships under specific environmental conditions may generate more insight than a global relationship can offer.

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 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.

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.

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.

AbstractRestoration in Mediterranean-climate grasslands is strongly impeded by lack of native propagules and competition with exotic grasses and forbs. We report on a study testing several methods for exotic plant control combined with planting native grasses to restore prairies in former agricultural land in coastal California. Specifically we compared tarping (shading out recently germinated seedlings with black plastic) once, tarping twice, topsoil removal, herbicide (glyphosate), and a control treatment in factorial combinations with or without wood mulch. Into each treatment we planted three native grass species (Elymus glaucus, Hordeum brachyantherum, andStipa pulchra) and monitored plant survival and cover for three growing seasons. Survival of native grass species was high in all treatments, but was slightly lower in unmulched soil removal and control treatments in the first 2 yr. Mulching, tarping, and herbicide were all effective in reducing exotic grass cover and enhancing native grass cover for the first 2 yr, but by the third growing season cover of the plant guilds and bare ground had mostly converged, primarily because of the declining effects of the initial treatments. Mulching and tarping were both considerably more expensive than herbicide treatment. Topsoil removal was less effective in increasing native grass cover likely because soil removal altered the surface hydrology in this system. Our results show that several treatments were effective in enhancing native grass establishment, but that longer term monitoring is needed to evaluate the efficacy of restoration efforts. The most appropriate approach to controlling exotics to restore specific grassland sites will depend not only on the effectiveness, but also on relative costs and site constraints.

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.

Enrichment planting in naturally recovering secondary forests or in tree plantations is increasingly being used as strategy to restore later-successional, large-seeded tropical forest trees. We seeded two tree species (Otoba novogranatensis and Ruagea glabra) in three agricultural sites in Southern Costa Rica: abandoned pastures, eight to ten year old secondary forests and three year old tree plantations (containing two N-fixing of four total tree species). We measured micrometeorological conditions, soil water content, plant water potential, leaf area, foliar C and N, and photosynthesis to better understand mechanistic responses of seedlings to conditions in the different successional habitats. Micrometeorological conditions, soil water content, and plant water potential were generally similar across habitats. Certain aspects of leaves (such as Specific Leaf Area and foliar N content), and photosynthesis (e.g. quantum yield and electron transport rate) were highest in the plantations, intermediate in the secondary forests, and lowest in abandoned pastures. Enhanced rates of photosynthetic biochemistry (such as Vxmax and Jmax) and Photosystem II efficiency (e.g. thermal energy dissipation) occurred in leaves from the plantations compared to the abandoned pastures, which may be related to higher leaf %N content. Results suggest that foliar N may be of greater importance than soil water content and micrometeorological factors in driving differences in photosynthetic processes across planting habitats. Planting seeds of these two species in plantations containing three year old trees (including two N-fixing species) enhances certain aspects of their photosynthesis and growth, compared to seedlings in abandoned pastures with non-native grasses, and thus can help increase forest recovery on abandoned agricultural lands.

Large-scale ecosystem restoration projects seldom undergo comprehensive evaluation to determine project effectiveness. Consequently, there are missed opportunities for learning and strategy refinement. Before our study, monitoring information from California’s middle Sacramento River had not been synthesized, despite restoration having been ongoing since 1989. Our assessment was based on the development and application of 36 quantitative ecological indicators. These indicators were used to characterize the status of terrestrial and floodplain resources (e.g., flora and fauna), channel dynamics (e.g., planform, geomorphology), and the flow regime. Indicators were also associated with specific goal statements of the CALFED Ecosystem Restoration Program. A collective weight of evidence approach was used to assess restoration success. Our synthesis demonstrates good progress in the restoration of riparian habitats, birds and other wildlife, but not in restoration of streamflows and geomorphic processes. For example, from 1999 to 2007, there was a > 600% increase in forest patch core size, and a 43% increase in the area of the river bordered by natural habitat > 500 m wide. Species richness of landbirds and beetles increased at restoration sites, as did detections of bats. However, degraded post-Shasta Dam streamflow conditions continued. Relative to pre-dam conditions, the average number of years that pass between flows that are sufficient to mobilize the bed, and those that are of sufficient magnitude to inundate the floodplain, increased by over 100%. Trends in geomorphic processes were strongly negative, with increases in the amount of bank hardened with riprap, and decreases in the area of floodplain reworked. Overall the channel simplified, becoming less sinuous with reduced overall channel length. Our progress assessment presents a compelling case for what needs to be done to further advance the ecological restoration of the river. The most important actions to be taken relate to promoting river meander and floodplain connectivity, and restoring components of the natural flow regime.

Restorationists commonly plant overstory and understory species simultaneously at the outset of restoration, but a mature forest canopy may be necessary to facilitate survival of light-intolerant understory species. We conducted two experiments in riparian forest restoration sites along the Sacramento River to determine whether: (1) introducing understory species is more successful at the beginning of restoration or after the canopy has developed; (2) canopy cover directly (via reduced light) or indirectly (by reducing non-native competition) facilitates survival of native understory species; and (3) seeding or planting seedlings of understory species is most effective. Seven native understory species were introduced as both seeds and seedlings into an experiment that manipulated canopy cover (open or canopy) and non-native grass competition (control or grass-specific herbicide). We conducted another experiment using shade cloth to directly test the effect of different light levels on seedling survival and growth of three species. Both experiments showed that four species (Aristolochia californica, Carex barbarae, Clematis ligusticifolia, and Vitis californica) had higher survival under low-light conditions (canopy or shade cloth). In contrast, three species (Artemisia douglasiana, Euthamia occidentalis and Rubus californica) had similar survival across open and canopy conditions. Cover of unplanted understory vegetation (mostly non-native) was much lower under the canopy than in open plots treated with grass-specific herbicide. Establishment from seed was generally low and highly variable. Our results suggest that to restore understory species in riparian forests in north–central California: light-intolerant understory species should be planted after canopy closure; canopy cover is more effective than grass-specific herbicide at reducing non-native understory cover; and planting seedlings is more successful than direct seeding.