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.

Abstract. We compiled 46 broadscale data sets of species richness for a wide range of terrestrial plant, invertebrate, and ectothermic vertebrate groups in all parts of the world to test the ability of metabolic theory to account for observed diversity gradients. The theory makes two related predictions: (1) ln-transformed richness is linearly associated with a linear, inverse transformation of annual temperature, and (2) the slope of the relationship is near À0.65. Of the 46 data sets, 14 had no significant relationship; of the remaining 32, nine were linear, meeting prediction 1. Model I (ordinary least squares, OLS) and model II (reduced major axis, RMA) regressions then tested the linear slopes against prediction 2. In the 23 data sets having nonlinear relationships between richness and temperature, split-line regression divided the data into linear components, and regressions were done on each component to test prediction 2 for subsets of the data. Of the 46 data sets analyzed in their entirety using OLS regression, one was consistent with metabolic theory (meeting both predictions), and one was possibly consistent. Using RMA regression, no data sets were consistent. Of 67 analyses of prediction 2 using OLS regression on all linear data sets and subsets, two were consistent with the prediction, and four were possibly consistent. Using RMA regression, one was consistent (albeit weakly), and four were possibly consistent. We also found that the relationship between richness and temperature is both taxonomically and geographically conditional, and there is no evidence for a universal response of diversity to temperature. Meta-analyses confirmed significant heterogeneity in slopes among data sets, and the combined slopes across studies were significantly lower than the range of slopes predicted by metabolic theory based on both OLS and RMA regressions. We conclude that metabolic theory, as currently formulated, is a poor predictor of observed diversity gradients in most terrestrial systems.

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.

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Ecosystem recovery from anthropogenic disturbances, either without human intervention or assisted by ecological restoration, is increasingly occurring worldwide. As ecosystems progress through recovery, it is important to estimate any resulting deficit in biodiversity and functions. Here we use data from 3,035 sampling plots worldwide, to quantify the interim reduction of biodiversity and functions occurring during the recovery process (that is, the ‘recovery debt'). Compared with reference levels, recovering ecosystems run annual deficits of 46–51% for organism abundance, 27–33% for species diversity, 32–42% for carbon cycling and 31–41% for nitrogen cycling. Our results are consistent across biomes but not across degrading factors. Our results suggest that recovering and restored ecosystems have less abundance, diversity and cycling of carbon and nitrogen than ‘undisturbed' ecosystems, and that even if complete recovery is reached, an interim recovery debt will accumulate. Under such circumstances, increasing the quantity of less-functional ecosystems through ecological restoration and offsetting are inadequate alternatives to ecosystem protection.

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Studies suggest that populations of different species do not decline equally after habitat loss. However, empirical tests have been confined to fine spatiotemporal scales and have rarely included plants. Using data from 89,365 forest survey plots covering peninsular Spain, we explored, for each of 34 common tree species, the relationship between probability of occurrence and the local cover of remaining forest. Twenty-four species showed a significant negative response to forest loss, so that decreased forest cover had a negative effect on tree diversity, but the responses of individual species were highly variable. Animal-dispersed species were less vulnerable to forest loss, with six showing positive responses to decreased forest cover. The results imply that plant-animal interactions help prevent the collapse of forest communities that suffer habitat destruction.

The length of time land has been available for colonization by plants and other organisms could provide a partial explanation of the contemporary richness gradients of trees. According to this hypothesis, increasing times of land availability entail higher chances of recolonization, which eventually have positive effects on tree richness. To test this, we generated a dataset of the Holarctic trees and evaluated the influence of cell age, a measure of the time since an area became free of ice, on the observed tree richness gradients. We found that cell age is associated with richness in both Europe and North America, after controlling for contemporary climate patterns, suggesting that the historical pattern of glacial retreat in response to post-Pleistocene global warming has left a signal still detectable after at least 14 000 yr. The results were consistent using a range of modelling approaches or whether Europe and North America were analyzed separately or in concert. We conclude that, although secondary to contemporary climate, the post-glacial recolonization hypothesis is broadly supported at temperate latitudes.