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Aim: Large-scale patterns linking energy availability, biological productivity and diversity form a central focus of ecology. Despite evidence that the activity and abundance of animals may be limited by climatic variables associated with regional biological productivity (e.g. mean annual precipitation and annual actual evapotranspiration), it is unclear whether plant-granivore interactions are themselves influenced by these climatic factors across broad spatial extents. We evaluated whether climatic conditions that are known to alter the abundance and activity of granivorous animals also affect rates of seed removal. Location: Eleven sites across temperate North America. Methods: We used a common protocol to assess the removal of the same seed species (Avena sativa) over a 2-day period. Model selection via the Akaike information criterion was used to determine a set of candidate binomial generalized linear mixed models that evaluated the relationship between local climatic data and post-dispersal seed predation. Results: Annual actual evapotranspiration was the single best predictor of the proportion of seeds removed. Annual actual evapotranspiration and mean annual precipitation were both positively related to mean seed removal and were included in four and three of the top five models, respectively. Annual temperature range was also positively related to seed removal and was an explanatory variable in three of the top four models. Main conclusions: Our work provides the first evidence that energy and precipitation, which are known to affect consumer abundance and activity, also translate to strong, predictable patterns of seed predation across a continent. More generally, these findings suggest that future changes in temperature and precipitation could have widespread consequences for plant species composition in grasslands, through impacts on plant recruitment.

Global change is associated with variable shifts in the annual production of aboveground plant biomass, suggesting localized sensitivities with unclear causal origins. Combining remotely sensed normalized difference vegetation index data since the 1980s with contemporary field data from 84 grasslands on 6 continents, we show a widening divergence in site-level biomass ranging from +51% to -34% globally. Biomass generally increased in warmer, wetter and species-rich sites with longer growing seasons and declined in species-poor arid areas. Phenological changes were widespread, revealing substantive transitions in grassland seasonal cycling. Grazing, nitrogen deposition and plant invasion were prevalent in some regions but did not predict overall trends. Grasslands are undergoing sizable changes in production, with implications for food security, biodiversity and carbon storage especially in arid regions where declines are accelerating., (© 2024. The Author(s), under exclusive licence to Springer Nature Limited.)

Scientists must have an integrative understanding of ecology and evolution across spatial and temporal scales to predict how species will respond to global change. Although comprehensively investigating these processes in nature is challenging, the infrastructure and data from long-term ecological research networks can support cross-disciplinary investigations. We propose using these networks to advance our understanding of fundamental evolutionary processes and responses to global change. For ecologists, we outline how long-term ecological experiments can be expanded for evolutionary inquiry, and for evolutionary biologists, we illustrate how observed long-term ecological patterns may motivate new evolutionary questions. We advocate for collaborative, multi-site investigations and discuss barriers to conducting evolutionary work at network sites. Ultimately, these networks offer valuable information and opportunities to improve predictions of species' responses to global change., (© 2024. Springer Nature Limited.)

Ecological restoration is critical for recovering degraded ecosystems but is challenged by variable success and low predictability. Understanding which outcomes are more predictable and less variable following restoration can improve restoration effectiveness. Recent theory asserts that the predictability of outcomes would follow an order from most to least predictable from coarse to fine community properties (physical structure > taxonomic diversity > functional composition > taxonomic composition) and that predictability would increase with more severe environmental conditions constraining species establishment. We tested this "hierarchy of predictability" hypothesis by synthesizing outcomes along an aridity gradient with 11 grassland restoration projects across the United States. We used 1829 vegetation monitoring plots from 227 restoration treatments, spread across 52 sites. We fit generalized linear mixed-effects models to predict six indicators of restoration outcomes as a function of restoration characteristics (i.e., seed mixes, disturbance, management actions, time since restoration) and used variance explained by models and model residuals as proxies for restoration predictability. We did not find consistent support for our hypotheses. Physical structure was among the most predictable outcomes when the response variable was relative abundance of grasses, but unpredictable for total canopy cover. Similarly, one dimension of taxonomic composition related to species identities was unpredictable, but another dimension of taxonomic composition indicating whether exotic or native species dominated the community was highly predictable. Taxonomic diversity (i.e., species richness) and functional composition (i.e., mean trait values) were intermittently predictable. Predictability also did not increase consistently with aridity. The dimension of taxonomic composition related to the identity of species in restored communities was more predictable (i.e., smaller residuals) in more arid sites, but functional composition was less predictable (i.e., larger residuals), and other outcomes showed no significant trend. Restoration outcomes were most predictable when they related to variation in dominant species, while those responding to rare species were harder to predict, indicating a potential role of scale in restoration predictability. Overall, our results highlight additional factors that might influence restoration predictability and add support to the importance of continuous monitoring and active management beyond one-time seed addition for successful grassland restoration in the United States., (© 2023 The Authors. Ecological Applications published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Premise: In 1879, Dr. William Beal buried 20 glass bottles filled with seeds and sand at a single site at Michigan State University. The goal of the experiment was to understand seed longevity in the soil, a topic of general importance in ecology, restoration, conservation, and agriculture, by periodically assaying germinability of these seeds over 100 years. The interval between germination assays has been extended and the experiment will now end after 221 years, in 2100., Methods: We dug up the 16th bottle in April 2021 and attempted to germinate the 141-year-old seeds it contained. We grew germinants to maturity and identified these to species by vegetative and reproductive phenotypes. For the first time in the history of this experiment, genomic DNA was sequenced to confirm species identities., Results: Twenty seeds germinated over the 244-day assay. Eight germinated in the first 11 days. All 20 belonged to the Verbascum genus: Nineteen were V. blattaria according to phenotype and ITS2 genotype; and one had a hybrid V. blattaria × V. thapsus phenotype and ITS2 genotype. In total, 20/50 (40%) of the original Verbascum seeds in the bottle germinated in year 141., Conclusions: While most species in the Beal experiment lost all seed viability in the first 60 years, a high percentage of Verbascum seeds can still germinate after 141 years in the soil. Long-term experiments such as this one are rare and invaluable for studying seed viability in natural soil conditions., (© 2023 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Although plant-soil feedbacks (interactions between plants and soils, often mediated by soil microbes, abbreviated as PSFs) are widely known to influence patterns of plant diversity at local and landscape scales, these interactions are rarely examined in the context of important environmental factors. Resolving the roles of environmental factors is important because the environmental context may alter PSF patterns by modifying the strength or even direction of PSFs for certain species. One important environmental factor that is increasing in scale and frequency with climate change is fire, though the influence of fire on PSFs remains essentially unexamined. By changing microbial community composition, fire may alter the microbes available to colonize the roots of plants and thus seedling growth post-fire. This has potential to change the strength and/or direction of PSFs, depending on how such changes in microbial community composition occur and the plant species with which the microbes interact. We examined how a recent fire altered PSFs of two leguminous, nitrogen-fixing tree species in Hawai'i. For both species, growing in conspecific soil resulted in higher plant performance (as measured by biomass production) than growing in heterospecific soil. This pattern was mediated by nodule formation, an important process for growth for legume species. Fire weakened PSFs for these species and therefore pairwise PSFs, which were significant in unburned soils, but were nonsignificant in burned soils. Theory suggests that positive PSFs such as those found in unburned sites would reinforce the dominance of species where they are locally dominant. The change in pairwise PSFs with burn status shows PSF-mediated dominance might diminish after fire. Our results demonstrate that fire can modify PSFs by weakening the legume-rhizobia symbiosis, which may alter local competitive dynamics between two canopy dominant tree species. These findings illustrate the importance of considering environmental context when evaluating the role of PSFs for plants., (© 2023 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America. This article has been contributed to by U.S. Government employees and their work is in the public domain in the USA.)

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions., (© 2023. The Author(s).)

Ecological restoration is essential for maintaining biodiversity in the face of dynamic, global changes in climate, human land use, and disturbance regimes. Effective restoration requires understanding bottlenecks in plant community recovery that exist today, while recognizing that these bottlenecks may relate to complex histories of environmental change. Such understanding has been a challenge because few long-term, well-replicated experiments exist to decipher the demographic processes influencing recovery for numerous species against the backdrop of multiyear variation in climate and management. We address this challenge through a long-term and geographically expansive experiment in longleaf pine savannas, an imperiled ecosystem and biodiversity hotspot in the southeastern United States. Using 48 sites at three locations spanning 480 km, the 8-y experiment manipulated initial seed arrival for 24 herbaceous plant species and presence of competitors to evaluate the impacts of climate variability and management actions (e.g., prescribed burning) on plant establishment and persistence. Adding seeds increased plant establishment of many species. Cool and wet climatic conditions, low tree density, and reduced litter depth also promoted establishment. Once established, most species persisted for the duration of the 8-y experiment. Plant traits were most predictive when tightly coupled to the process of establishment. Our results illustrate how seed additions can restore plant diversity and how interannual climatic variation affects the dynamics of plant communities across a large region. The significant effects of temperature and precipitation inform how future climate may affect restoration and conservation via large-scale changes in the fundamental processes of establishment and persistence.

Relationships between biodiversity and ecosystem functioning depend on the processes structuring community assembly. However, predicting biodiversity-ecosystem functioning (BEF) relationships based on community assembly remains challenging because assembly outcomes are often contingent on history and the consequences of history for ecosystem functions are poorly understood. In a grassland restoration experiment, we isolated the role of history for the relationships between plant biodiversity and multiple ecosystem functions by initiating assembly in three different years, while controlling for all other aspects of community assembly. We found that two aspects of assembly history-establishment year and succession-altered species and trait community trajectories, which in turn altered net primary productivity, decomposition rates, and floral resources. Moreover, history altered BEF relationships (which ranged from positive to negative), both within and across functions, by modifying the causal pathways linking species identity, traits, diversity, and ecosystem functions. Our results show that the interplay of deterministic succession and environmental stochasticity during establishment mediate historical contingencies that cause variation in biodiversity and ecosystem functions, even under otherwise identical assembly conditions. An explicit attention to history is needed to understand why biodiversity-ecosystem function relationships vary in natural ecosystems: a critical question at the intersection of fundamental theory and applications to environmental change biology and ecosystem restoration., (© 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.)

Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species' evolutionary history of grazing continues to shape the response of the world's grasslands to changing mammalian herbivory., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Ecological restoration projects often have variable and unpredictable outcomes, and these can limit the overall impact on biodiversity. Previous syntheses have investigated restoration effectiveness by comparing average restored conditions to average conditions in unrestored or reference systems. Here, we provide the first quantification of the extent to which restoration affects both the mean and variability of biodiversity outcomes, through a global meta-analysis of 83 terrestrial restoration studies. We found that, relative to unrestored (degraded) sites, restoration actions increased biodiversity by an average of 20%, while decreasing the variability of biodiversity (quantified by the coefficient of variation) by an average of 14%. As restorations aged, mean biodiversity increased and variability decreased relative to unrestored sites. However, restoration sites remained, on average, 13% below the biodiversity of reference (target) ecosystems, and were characterised by higher (20%) variability. The lower mean and higher variability in biodiversity at restored sites relative to reference sites remained consistent over time, suggesting that sources of variation (e.g. prior land use, restoration practices) have an enduring influence on restoration outcomes. Our results point to the need for new research confronting the causes of variability in restoration outcomes, and close variability and biodiversity gaps between restored and reference conditions., (© 2022 The Authors. Ecology Letters published by John Wiley & Sons Ltd.)

Habitat loss and fragmentation are leading causes of species declines, driven in part by reduced dispersal. Isolating the effects of fragmentation on dispersal, however, is daunting because the consequences of fragmentation are typically intertwined, such as reduced connectivity and increased prevalence of edge effects. We used a large-scale landscape experiment to separate consequences of fragmentation on seed dispersal, considering both distance and direction of local dispersal. We evaluated seed dispersal for five wind- or gravity-dispersed, herbaceous plant species that were planted at different distances from habitat edges, within fragments that varied in their connectivity and shape (edge-to-area ratio). Dispersal distance was affected by proximity and direction relative to the nearest edge. For four of five species, dispersal distances were greater further from habitat edges and when seeds dispersed in the direction of the nearest edge. Connectivity and patch edge-to-area ratio had minimal effects on local dispersal. Our findings illustrate how some, but not all, landscape changes associated with fragmentation can affect the key population process of seed dispersal., (© 2021 The Ecological Society of America.)

Restoration in this era of climate change comes with a new challenge: anticipating how best to restore populations to persist under future climate conditions. Specifically, it remains unknown whether locally adapted or warm-adapted seeds best promote native plant community restoration in the warmer conditions predicted in the future and whether local or warm-adapted soil microbial communities could mitigate plant responses to warming. This may be especially relevant for biomes spanning large climatic gradients, such as the North American tallgrass prairie. Here, we used a short-term mesocosm experiment to evaluate how seed provenances (Local Northern region, Non-local Northern region, Non-local Southern region) of 10 native tallgrass prairie plants (four forbs, two legumes, and four grasses) responded to warmer conditions predicted in the future and how soil microbial communities from those three regions influenced these responses. Warming and seed provenance affected plant community composition and warming decreased plant diversity for all three seed provenances. Plant species varied in their individual responses to warming, and across species, we detected no consistent differences among the three provenances in terms of biomass response to warming and few strong effects of soil provenance. Our work provides evidence that warming, in part, may reduce plant diversity and affect restored prairie composition. Because the southern provenance did not consistently outperform others under warming and we found little support for the "local is best" paradigm currently dominating restoration practice, identifying appropriate seed provenances to promote restoration success both now and in future warmer environments may be challenging. Due to the idiosyncratic responses across species, we recommend that land managers compare seeds from different regions for each species to determine which seed provenance performs best under warming and in restoration for tallgrass prairies., (© 2021 by the Ecological Society of America.)

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5-12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone., (© 2021 by the Ecological Society of America.)

Ecological restoration is a global priority, with potential to reverse biodiversity declines and promote ecosystem functioning. Yet, successful restoration is challenged by lingering legacies of past land-use activities, which are pervasive on lands available for restoration. Although legacies can persist for centuries following cessation of human land uses such as agriculture, we currently lack understanding of how land-use legacies affect entire ecosystems, how they influence restoration outcomes, or whether restoration can mitigate legacy effects. Using a large-scale experiment, we evaluated how restoration by tree thinning and land-use legacies from prior cultivation and subsequent conversion to pine plantations affect fire-suppressed longleaf pine savannas. We evaluated 45 ecological properties across four categories: 1) abiotic attributes, 2) organism abundances, 3) species diversity, and 4) species interactions. The effects of restoration and land-use legacies were pervasive, shaping all categories of properties, with restoration effects roughly twice the magnitude of legacy effects. Restoration effects were of comparable magnitude in savannas with and without a history of intensive human land use; however, restoration did not mitigate numerous legacy effects present prior to restoration. As a result, savannas with a history of intensive human land use supported altered properties, especially related to soils, even after restoration. The signature of past human land-use activities can be remarkably persistent in the face of intensive restoration, influencing the outcome of restoration across diverse ecological properties. Understanding and mitigating land-use legacies will maximize the potential to restore degraded ecosystems., Competing Interests: The authors declare no competing interest.

The species pool concept has advanced our understanding for how biodiversity is coupled at local and regional scales. However, it remains unclear how species pool size, the number of species available to disperse to a site, influences community assembly across spatial scales. We provide one of the first studies that assesses diversity across scales after experimentally assembling grassland communities from species pools of different sizes. We show that species pool size causes scale-dependent effects on diversity in grasslands undergoing restoration by altering the shape of the species-area relationship (SAR). Specifically, larger species pools increased the slope of the SAR, but not the intercept, suggesting that dispersal from a larger pool causes species to be more spatially aggregated. This increased aggregation appears to be caused by sampling effects due to fewer individuals arriving per species, rather than stronger species sorting across variation in soil moisture. These scale-dependent effects suggest that studies evaluating species pools at a single, small scale may underestimate their effects, thereby contributing to uncertainty about the importance of regional processes for community assembly and their consequences for ecological restoration., (© 2020 by the Ecological Society of America.)

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities.

Ecological restoration - the rebuilding of damaged or destroyed ecosystems - is a critical component of conservation efforts, but is hindered by inconsistent, unpredictable outcomes. We investigated a source of this variation that is anecdotally suggested by practitioners, but for which empirical evidence is rare: the weather conditions during the first growing season after planting. The idea of whether natural communities face long-term consequences from conditions even many years in the past, called historical contingency, is a debated idea in ecological research. Using a large dataset (83 sites) across a wide geographic distribution (three states), we find evidence that precipitation and temperatures in the planting year (2-19 years before present) affected the relative dominance of the sown (native target species) and non-sown (mostly non-native) species. We find strong support for lasting planting year weather effects in restored tallgrass prairies, thereby supporting the historically contingent model of community assembly in a real-world setting.

Establishment and persistence are central to community assembly and are determined by how traits interact with the environment to determine performance (trait-environment interactions). Community assembly studies have rarely considered such trait-environment interactions, however, which can lead to incorrect inferences about how traits affect assembly. We evaluated how functional traits, environmental conditions, and trait-environment interactions structure plant establishment, as a measure of performance. Within 12 prairie restorations created by sowing 70 species, we quantified environmental conditions and counted individuals of each seeded species to quantify first-year establishment. Three trait-environment interactions structured establishment. Leaf nitrogen interacted with herbivore pressure, as low leaf nitrogen species established relatively better under higher herbivory than species with high leaf nitrogen. Soil moisture interacted with root mass fraction (RMF), with low-RMF species establishing better with low soil moisture and higher-RMF species better on wetter soils. Specific leaf area (SLA) interacted with light availability, as low-SLA species established better under high light conditions and high-SLA species under low light conditions. Our work illustrates how community assembly can be better described by trait-environment interactions than correlating traits or environment with performance. This knowledge can assist species selection to maximize restoration success., (© 2020 by the Ecological Society of America.)

Deleterious effects of habitat fragmentation and benefits of connecting fragments could be significantly underestimated because changes in colonization and extinction rates that drive changes in biodiversity can take decades to accrue. In a large and well-replicated habitat fragmentation experiment, we find that annual colonization rates for 239 plant species in connected fragments are 5% higher and annual extinction rates 2% lower than in unconnected fragments. This has resulted in a steady, nonasymptotic increase in diversity, with nearly 14% more species in connected fragments after almost two decades. Our results show that the full biodiversity value of connectivity is much greater than previously estimated, cannot be effectively evaluated at short time scales, and can be maximized by connecting habitat sooner rather than later., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Agricultural land use is a leading cause of habitat degradation, leaving a legacy of ecological impacts long after agriculture has ceased. Yet the mechanisms for legacy effects, such as altered plant community composition, are not well understood. In particular, whether plant community recovery is limited by an inability of populations to establish within post-agricultural areas, owing to altered environmental conditions within these areas, remains poorly known. We evaluated this hypothesis of post-agricultural establishment limitation through a field experiment within longleaf pine woodlands in South Carolina (USA) and a greenhouse experiment using field-collected soils from these sites. In the field, we sowed seeds of 12 understory plant species associated with remnants (no known history of agriculture) into 27 paired remnant and post-agricultural woodlands. We found that post-agricultural woodlands supported higher establishment, resulting in greater species richness of sown species. These results were context dependent, however, with higher establishment in post-agricultural woodlands only when sites were frequently burned, had less leaf litter, or had less sandy soils. In the greenhouse, we found that agricultural history had no impact on plant growth or survival, suggesting that establishment limitation is unlikely driven by differences in soils associated with agricultural history when environmental conditions are not stressful. Rather, the potential for establishment in post-agricultural habitats can be higher than in remnant habitats, with the strength of this effect determined by fire frequency and soil characteristics.

There is strong evidence for a positive relationship between biodiversity and ecosystem functioning at local spatial scales. However, how different aspects of biodiversity relate to multiple ecosystem functions (multifunctionality) across heterogeneous landscapes, and how the magnitude of biodiversity, dominant species, and environmental effects on functioning compare, remain poorly understood. We compared relationships between plant phylogenetic, functional, and taxonomic diversity and ecosystem multifunctionality across 29 restored grasslands. Functional diversity was positively associated with multifunctionality, more strongly than other diversity measures; however, landscape composition explained nearly four times more variation in multifunctionality than did functional diversity, with plots within human-modified landscapes supporting lower multifunctionality. Individual functions were typically more strongly correlated with environmental variables than with diversity. We also found that abundance of the dominant species, Andropogon gerardii, was positively correlated with multifunctionality. Plant diversity, dominant species, and underlying environmental conditions underpin ecosystem multifunctionality in grasslands, but how biodiversity is measured matters for the strength and direction of biodiversity-ecosystem function relationships. Finally, in natural systems environmental variation unrelated to local biodiversity is important for determining ecosystem functioning., (© 2019 by the Ecological Society of America.)

Conservation and restoration projects often involve starting new populations by introducing individuals into portions of their native or projected range. Such efforts can help meet many related goals, including habitat creation, ecosystem service provisioning, assisted migration, and the reintroduction of imperiled species following local extirpation. The outcomes of reintroduction efforts, however, are highly variable, with results ranging from local extinction to dramatic population growth; reasons for this variation remain unclear. Here, we ask whether population growth following plant reintroductions is governed by variation at two scales: the scale of individual habitat patches to which individuals are reintroduced, and larger among-landscape scales in which similar patches may be situated in landscapes that differ in matrix type, soil conditions, and other factors. Quantifying demographic variation at these two scales will help prioritize locations for introduction and, once introductions take place, forecast population growth. This work took place within a large-scale habitat fragmentation experiment, where individuals of two perennial forb species were reintroduced into eight replicate ~50-ha landscapes, each containing a set of five ~1-ha patches that varied in their degree of isolation (connected by habitat corridors or unconnected) and edge-to-area ratio. Using data on individual growth, survival, reproductive output, and recruitment collected one to two years after reintroduction, we developed models to forecast population growth, then compared forecasts to observed population sizes, three and six years later. Both the type of patch (connected and unconnected) and identity of the landscape to which individuals were reintroduced had effects on forecasted population growth rates, but only variation associated with landscape identity was an accurate predictor of subsequently observed population growth rates. Models that did not include landscape identity had minimal forecasting ability, revealing the key importance of variation at this scale for accurate prediction. Of the five demographic rates used to model population dynamics, seed production was the most important source of forecast error in population growth rates. Our results point to the importance of accounting for landscape-scale variation in demographic models and demonstrate how such models might assist with prioritizing particular landscapes for species reintroduction projects., (© 2019 by the Ecological Society of America.)

The loss of biodiversity at local and larger scales has potentially dramatic effects on ecosystem functioning. Many studies have shown that ecosystem functioning depends on biodiversity, but the role of beta diversity, spatial variation in community composition, is less clear than that of local-scale (alpha) diversity. To test the hypothesis that beta diversity would increase ecosystem multifunctionality through variation in species functional traits, we gathered data on plant community composition, plant functional traits, and seven ecosystem functions across 29 restored prairies. We found that averaged multifunctionality (mean of seven ecosystem functions) increased with both taxonomic beta diversity and functional beta diversity. The abundance of the dominant species, big bluestem, played a more minor role, suggesting a limited role for the selection effect. Neither taxonomic nor functional alpha richness was associated with multifunctionality, though this finding may be sensitive to the identity of the functions included because alpha diversity was associated with some individual functions in opposing directions. These findings suggest that in systems structured largely by natural processes, beta diversity (a patchwork of functionally different plant communities) and dominant species abundance may be more important than alpha diversity in fostering ecosystem multifunctionality. These findings suggest the need for an increased focus on community heterogeneity to reestablish functional ecosystems during restoration.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experiments show that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of community composition and the potential interaction between diversity at different scales in maintaining ecosystem functioning remains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis of eight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands-those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)-had higher levels of multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities.

Habitat fragmentation affects species and their interactions through intertwined mechanisms that include changes to fragment area, shape, connectivity and distance to edge. Disentangling these pathways is a fundamental challenge of landscape ecology and will help identify ecological processes important for management of rare species or restoration of fragmented habitats. In a landscape experiment that manipulated connectivity, fragment shape, and distance to edge while holding fragment area constant, we examined how fragmentation impacts herbivory and growth of nine plant species in longleaf pine savanna. Probability of herbivory in open habitat was strongly dependent on proximity to forest edge for every species, increasing with distance to edge in six species (primarily grasses and annual forbs) and decreasing in three species (perennial forbs and a shrub). In the two species of perennial forbs, these edge effects were dependent on fragment shape; herbivory strongly decreased with distance to edge in fragments of two shapes, but not in a third shape. For most species, however, probability of herbivory was unrelated to connectivity or fragment shape. Growth was generally determined more strongly by leaf herbivory than by distance to edge, fragment shape, or connectivity. Taken together, these results demonstrate consistently strong edge effects on herbivory, one of the most important biotic factors determining plant growth and demography. Our results contrast with the generally inconsistent results of observational studies, likely because our experimental approach enabled us to tease apart landscape processes that are typically confounded., (© 2016 by the Ecological Society of America.)

Intensive land use activities, such as agriculture, are a leading cause of biodiversity loss and can have lasting impacts on ecological systems. Yet, few studies have investigated how land-use legacies impact phylogenetic diversity (the total amount of evolutionary history in a community) or how restoration activities might mitigate legacy effects on biodiversity. We studied ground-layer plant communities in 27 pairs of Remnant (no agricultural history) and Post-agricultural (agriculture abandoned >60 yr ago) longleaf pine savannas, half of which we restored by thinning trees to reinstate open savanna conditions. We found that agricultural history had no impact on species richness, but did alter community composition and reduce phylogenetic diversity by 566 million years/1,000 m 2 . This loss of phylogenetic diversity in post-agricultural savannas was due to, in part, a reduction in the average evolutionary distance between pairs of closely related species, that is, increased phylogenetic clustering. Habitat restoration increased species richness by 27% and phylogenetic diversity by 914 million years but did not eliminate the effects of agricultural land use on community composition and phylogenetic structure. These results demonstrate the persistence of agricultural legacies, even in the face of intensive restoration efforts, and the importance of considering biodiversity broadly when evaluating human impacts on ecosystems., (© 2016 by the Ecological Society of America.)

Habitat fragmentation can create significant impediments to dispersal. A technique to increase dispersal between otherwise isolated fragments is the use of corridors. Although previous studies have compared dispersal between connected fragments to dispersal between unconnected fragments, it remains unknown how dispersal between fragments connected by a corridor compares to dispersal in unfragmented landscapes. To assess the extent to which corridors can restore dispersal in fragmented landscapes to levels observed in unfragmented landscapes, we employed a stable-isotope marking technique to track seeds within four unfragmented landscapes and eight experimental landscapes with fragments connected by corridors. We studied two wind- and two bird-dispersed plant species, because previous community-based research showed that dispersal mode explains how connectivity effects vary among species. We constructed dispersal kernels for these species in unfragmented landscapes and connected fragments by marking seeds in the center of each landscape with 'IN and then recovering marked seeds in seed traps at distances up to 200 m. For the two wind-dispersed plants, seed dispersal kernels were similar in unfragmented landscapes and connected fragments. In contrast, dispersal kernels of bird-dispersed seeds were both affected by fragmentation and differed in the direction of the impact: Morella cerifera experienced more and Rhus copallina experienced less long-distance dispersal in unfragmented than in connected landscapes. These results show that corridors can facilitate dispersal probabilities comparable to those observed in unfragmented landscapes. Although dispersal mode may provide useful broad predictions, we acknowledge that similar species may respond uniquely due to factors such as seasonality and disperser behavior. Our results further indicate that prior work has likely underestimated dispersal distances of wind-dispersed plants and that factors altering long-distance dispersal may have a greater impact on the spread of species than previously thought.

Despite broad recognition that habitat loss represents the greatest threat to the world's biodiyersity, a mechanistic understanding of how habitat loss and associated fragmentation affect ecological systems has proven remarkably challenging. The challenge stems from the multiple interdependent ways that landscapes change following fragmentation and the ensuing complex impacts on populations and communities of interacting species. We confronted these challenges by evaluating how fragmentation affects individual plants through interactions with animals, across five herbaceous species native to longleaf pine savannas. We created a replicated landscape experiment that provides controlled tests of three major fragmentation effects (patch isolation, patch shape [i.e., edge-to-area ratio], and distance to edge), established experimental founder populations of the five species to control for spatial distributions and densities of individual plants, and employed structural equation modeling to evaluate the effects of fragmentation on plant reproductive output and the degree to which these impacts are mediated through altered herbivory, pollination, or pre-dispersal seed predation. Across species, the most consistent response to fragmentation was a reduction in herbivory. Herbivory, however, had little impact.on plant reproductive output, and thus we found little evidence for any resulting benefit to plants in fragments. In contrast, fragmentation rarely impacted pollination or pre-dispersal seed predation, but both of these interactions had strong and consistent impacts on plant reproductive output. As a result, our models robustly predicted plant reproductive output (r2 = 0.52-0.70), yet due to the weak effects of fragmentation on pollination and pre-dispersal seed predation, coupled with the weak effect of herbivory on plant reproduction, the effects of fragmentation on reproductive output were generally small in magnitude and inconsistent. This work provides mechanistic insight into landscape-scale variation in plant reproductive success, the relative importance of plant-animal interactions for structuring these dynamics, and the nuanced nature of how habitat fragmentation can affect populations and communities of interacting species.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

We conducted an analysis of global forest cover to reveal that 70% of remaining forest is within 1 km of the forest's edge, subject to the degrading effects of fragmentation. A synthesis of fragmentation experiments spanning multiple biomes and scales, five continents, and 35 years demonstrates that habitat fragmentation reduces biodiversity by 13 to 75% and impairs key ecosystem functions by decreasing biomass and altering nutrient cycles. Effects are greatest in the smallest and most isolated fragments, and they magnify with the passage of time. These findings indicate an urgent need for conservation and restoration measures to improve landscape connectivity, which will reduce extinction rates and help maintain ecosystem services.

Land-use legacies are known to shape the diversity and distribution of plant communities, but we lack an understanding of whether historical land use influences community responses to contemporary disturbances. Because human-modified landscapes often bear a history of multiple land-use activities, this contingency can challenge our understanding of land-use impacts on plant diversity. We address this contingency by evaluating how beta diversity (the spatial variability of species composition), an important component of regional biodiversity, is shaped by interactions between historical agriculture and prescribed fire, two prominent disturbances that are often coincident in terrestrial ecosystems. At three study locations spanning 450 km in the southeastern United States, we surveyed longleaf pine woodland understory plant communities across 232 remnant and post-agricultural sites with differing prescribed fire regimes. Our results demonstrate that agricultural legacies are a strong predictor of beta diversity, but the direction of this land-use effect differed among the three study locations. Further, although beta diversity increased with prescribed fire frequency at each study location, this effect was influenced by agricultural land-use history, such that positive fire effects were only documented among sites that lacked a history of agriculture at two of our three study locations. Our study not only highlights the role of historical agriculture in shaping beta diversity in a fire-maintained ecosystem but also illustrates how this effect can be contingent upon fire regime and geographic location. We suggest that interactions among historical and contemporary land-use activities may help to explain dissimilarities in plant communities among sites in human-dominated landscapes.

Despite many studies showing that landscape corridors increase dispersal and species richness for disparate taxa, concerns persist that corridors can have unintended negative effects. In particular, some of the same mechanisms that underlie positive effects of corridors on species of conservation interest may also increase the spread and impact of antagonistic species (e.g., predators and pathogens), foster negative effects of edges, increase invasion by exotic species, increase the spread of unwanted disturbances such as fire, or increase population synchrony and thus reduce persistence. We conducted a literature review and meta-analysis to evaluate the prevalence of each of these negative effects. We found no evidence that corridors increase unwanted disturbance or non-native species invasion; however, these have not been well-studied concerns (1 and 6 studies, respectively). Other effects of corridors were more often studied and yielded inconsistent results; mean effect sizes were indistinguishable from zero. The effect of edges on abundances of target species was as likely to be positive as negative. Corridors were as likely to have no effect on antagonists or population synchrony as they were to increase those negative effects. We found 3 deficiencies in the literature. First, despite studies on how corridors affect predators, there are few studies of related consequences for prey population size and persistence. Second, properly designed studies of negative corridor effects are needed in natural corridors at scales larger than those achievable in experimental systems. Third, studies are needed to test more targeted hypotheses about when corridor-mediated effects on invasive species or disturbance may be negative for species of management concern. Overall, we found no overarching support for concerns that construction and maintenance of habitat corridors may result in unintended negative consequences. Negative edge effects may be mitigated by widening corridors or softening edges between corridors and the matrix. Other negative effects are relatively small and manageable compared with the large positive effects of facilitating dispersal and increasing diversity of native species., (© 2014 Society for Conservation Biology.)

Landscape corridors are commonly used to mitigate negative effects of habitat fragmentation, but concerns persist that they may facilitate the spread of invasive species. In a replicated landscape experiment of open habitat, we measured effects of corridors on the invasive fire ant, Solenopsis invicta, and native ants. Fire ants have two social forms: polygyne, which tend to disperse poorly but establish at high densities, and monogyne, which disperse widely but establish at lower densities. In landscapes dominated by polygyne fire ants, fire ant abundance was higher and native ant diversity was lower in habitat patches connected by corridors than in unconnected patches. Conversely, in landscapes dominated by monogyne fire ants, connectivity had no influence on fire ant abundance and native ant diversity. Polygyne fire ants dominated recently created landscapes, suggesting that these corridor effects may be transient. Our results suggest that corridors can facilitate invasion and they highlight the importance of considering species' traits when assessing corridor utility.

Human alterations to nutrient cycles and herbivore communities are affecting global biodiversity dramatically. Ecological theory predicts these changes should be strongly counteractive: nutrient addition drives plant species loss through intensified competition for light, whereas herbivores prevent competitive exclusion by increasing ground-level light, particularly in productive systems. Here we use experimental data spanning a globally relevant range of conditions to test the hypothesis that herbaceous plant species losses caused by eutrophication may be offset by increased light availability due to herbivory. This experiment, replicated in 40 grasslands on 6 continents, demonstrates that nutrients and herbivores can serve as counteracting forces to control local plant diversity through light limitation, independent of site productivity, soil nitrogen, herbivore type and climate. Nutrient addition consistently reduced local diversity through light limitation, and herbivory rescued diversity at sites where it alleviated light limitation. Thus, species loss from anthropogenic eutrophication can be ameliorated in grasslands where herbivory increases ground-level light.

Determining how widespread human-induced changes such as habitat loss, landscape fragmentation, and climate instability affect populations, communities, and ecosystems is one of the most pressing environmental challenges. Critical to this challenge is understanding how these changes are affecting the movement abilities and dispersal trajectories of organisms and what role conservation planning can play in promoting movement among remaining fragments of suitable habitat. Whereas evidence is mounting for how conservation strategies such as corridors impact animal movement, virtually nothing is known for species dispersed by wind, which are often mistakenly assumed to not be limited by dispersal. Here, we combine mechanistic dispersal models, wind measurements, and seed releases in a large-scale experimental landscape to show that habitat corridors affect wind dynamics and seed dispersal by redirecting and bellowing airflow and by increasing the likelihood of seed uplift. Wind direction interacts with landscape orientation to determine when corridors provide connectivity. Our results predict positive impacts of connectivity and patch shape on species richness of wind-dispersed plants, which we empirically illustrate using 12 y of data from our experimental landscapes. We conclude that habitat fragmentation and corridors strongly impact the movement of wind-dispersed species, which has community-level consequences.

Ecological restoration is frequently guided by reference conditions describing a successfully restored ecosystem; however, the causes and magnitude of ecosystem degradation vary, making simple knowledge of reference conditions insufficient for prioritizing and guiding restoration. Ecological reference models provide further guidance by quantifying reference conditions, as well as conditions at degraded states that deviate from reference conditions. Many reference models remain qualitative, however, limiting their utility. We quantified and evaluated a reference model for southeastern U.S. longleaf pine woodland understory plant communities. We used regression trees to classify 232 longleaf pine woodland sites at three locations along the Atlantic coastal plain based on relationships between understory plant community composition, soils (which broadly structure these communities), and factors associated with understory degradation, including fire frequency, agricultural history, and tree basal area. To understand the spatial generality of this model, we classified all sites together and for each of three study locations separately. Both the regional and location-specific models produced quantifiable degradation gradients-i.e., progressive deviation from conditions at 38 reference sites, based on understory species composition, diversity and total cover, litter depth, and other attributes. Regionally, fire suppression was the most important degrading factor, followed by agricultural history, but at individual locations, agricultural history or tree basal area was most important. At one location, the influence of a degrading factor depended on soil attributes. We suggest that our regional model can help prioritize longleaf pine woodland restoration across our study region; however, due to substantial landscape-to-landscape variation, local management decisions should take into account additional factors (e.g., soil attributes). Our study demonstrates the utility of quantifying degraded states and provides a series of hypotheses for future experimental restoration work. More broadly, our work provides a framework for developing and evaluating reference models that incorporate multiple, interactive anthropogenic drivers of ecosystem degradation.

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions., (© 2013 John Wiley & Sons Ltd.)

Although savanna trees and forest trees are thought to represent distinct functional groups with different effects on ecosystem processes, few empirical studies have examined these effects. In particular, it remains unclear if savanna and forest trees differ in their ability to coexist with understory plants, which comprise the majority of plant diversity in most savannas. We used structural equation modeling (SEM) and data from 157 sites across three locations in the southeastern United States to understand the effects of broadleaf savanna trees, broadleaf forest trees, and pine trees on savanna understory plant communities. After accounting for underlying gradients in fire frequency and soil moisture, abundances (i.e., basal area and stem density) of forest trees and pines, but not savanna trees, were negatively correlated with the cover and density (i.e., local-scale species richness) of C4 graminoid species, a defining savanna understory functional group that is linked to ecosystem flammability. In analyses of the full understory community, abundances of trees from all functional groups were negatively correlated with species density and cover. For both the C4 and full communities, fire frequency promoted understory plants directly, and indirectly by limiting forest tree abundance. There was little indirect influence of fire on the understory mediated through savanna trees and pines, which are morefire tolerant than forest trees. We conclude that tree functional identity is an important factor that influences overstory tree relationships with savanna understory plant communities. In particular, distinct relationships between trees and C4 graminoids have implications for grass-tree coexistence and vegetation-fire feedbacks that maintain savanna environments and their associated understory plant diversity.

Local species diversity is maintained over ecological time by a balance between dispersal and species interactions. Local-regional species richness relationships are often used to investigate the relative importance of these two processes and the scales at which they operate. For communities undergoing succession, theory predicts a temporal progression in local-regional species richness relationships: from no relationship to positive linear to saturating. However, observational tests have been mixed, and experiments have been rare. Using a replicated large-scale experiment, we evaluate the impact of two dispersal-governing processes at the regional scale, connectivity and shape of the region (i.e., patches), on the progression of local-regional species richness relationships for plant communities undergoing succession. Regional connectivity accelerates the transition from no relationship to a positive linear relationship, while the shape of the region has no consistent effect nine years post-disturbance. Our results experimentally demonstrate the importance of dispersal in structuring local-regional species richness relationships over time and suggest that conservation corridors among regions can increase local diversity through regional enrichment of plant communities undergoing reassembly.

Landscape corridors, strips of habitat that connect otherwise isolated habitat patches, are commonly employed during management of fragmented landscapes. To date, most reported effects of corridors have been positive; however, there are long-standing concerns that corridors may have unintended consequences. Here, we address concerns over whether corridors promote propagation of disturbances such as fire. We collected data during prescribed fires in the world's largest and best replicated corridor experiment (Savannah River Site, South Carolina, USA), six -50-ha landscapes of open (shrubby/herbaceous) habitat within a pine plantation matrix, to test several mechanisms for how corridors might influence fire. Corridors altered patterns of fire temperature through a direct connectivity effect and an indirect edge effect. The connectivity effect was independent of fuel levels and was consistent with a hypothesized wind-driven "bellows effect." Edges, a consequence of corridor implementation, elevated leaf litter (fuel) input from matrix pine trees, which in turn increased fire temperatures. We found no evidence for corridors or edges impacting patterns of fire spread: plots across all landscape positions burned with similar probability. Impacts of edges and connectivity on fire temperature led to changes in vegetation: hotter-burning plots supported higher bunch grass cover during the field season after burning, suggesting implications for woody/herbaceous species coexistence. To our knowledge, this represents the first experimental evidence that corridors can modify landscape-scale patterns of fire intensity. Corridor impacts on fire should be carefully considered during landscape management, both in the context of how corridors connect or break distributions of fuels and the desired role of fire as a disturbance, which may range from a management tool to an agent to be suppressed. In our focal ecosystem, longleaf pine woodland, corridors might provide a previously unrecognized benefit during prescribed burning activities, by promoting fire intensity, which may assist in promoting plant biodiversity.

Habitat corridors, a common management strategy for increasing connectivity in fragmented landscapes, have experimentally validated positive influences on species movement and diversity. However, long-standing concerns that corridors could negatively impact native species by spreading antagonists, such as disease, remain largely untested. Using a large-scale, replicated experiment, we evaluated whether corridors increase the incidence of plant parasites. We found that corridor impacts varied with parasite dispersal mode. Connectivity provided by corridors increased incidence of biotically dispersed parasites (galls on Solidago odora) but not of abiotically dispersed parasites (foliar fungi on S. odora and three Lespedeza spp.). Both biotically and abiotically dispersed parasites responded to edge effects, but the direction of responses varied across species. Although our results require additional tests for generality to other species and landscapes, they suggest that, when establishing conservation corridors, managers should focus on mitigating two potential negative effects: the indirect effects of narrow corridors in creating edges and direct effects of corridors in enhancing connectivity of biotically dispersed parasites.

The practice of ecological restoration is a primary option for increasing levels of biodiversity by modifying human-altered ecosystems. The scientific discipline of restoration ecology provides conceptual guidance and tests of restoration strategies, with the ultimate goal of predictive landscape restoration. I construct a conceptual model for restoration of biodiversity, based on site-level (e.g., biotic and abiotic) conditions, landscape (e.g, interpatch connectivity and patch geometry), and historical factors (e.g., species arrival order and land-use legacies). I then ask how well restoration ecology has addressed the various components of this model. During the past decade, restoration research has focused largely on how the restoration of site-level factors promotes species diversity-primarily of plants. Relatively little attention has been paid to how landscape or historical factors interplay with restoration, how restoration influences functional and genetic components of biodiversity, or how a suite of less-studied taxa might be restored. I suggest that the high level of variation seen in restoration outcomes might be explained, at least in part, by the contingencies placed on site-level restoration by landscape and historical factors and then present a number of avenues for future research to address these often ignored linkages in the biodiversity restoration model. Such work will require carefully conducted restoration experiments set across multiple sites and many years. It is my hope that by considering how space and time influence restoration, we might move restoration ecology in a direction of stronger prediction, conducted across landscapes, thus providing feasible restoration strategies that work at scales over which biodiversity conservation occurs.

Conservation efforts typically focus on maximizing biodiversity in protected areas. The space available for reserves is limited, however, and conservation efforts must increasingly consider how management of protected areas can promote biodiversity beyond reserve borders. Habitat corridors are considered an important feature of reserves because they facilitate movement of organisms between patches, thereby increasing species richness in those patches. Here we demonstrate that by increasing species richness inside target patches, corridors additionally benefit biodiversity in surrounding non-target habitat, a biodiversity "spillover" effect. Working in the world's largest corridor experiment, we show that increased richness extends for approximately 30% of the width of the 1-ha connected patches, resulting in 10-18% more vascular plant species around patches of target habitat connected by corridors than around unconnected but otherwise equivalent patches of habitat. Furthermore, corridor-enhanced spillover into non-target habitat can be predicted by a simple plant life-history trait: seed dispersal mode. Species richness of animal-dispersed plants in non-target habitat increased in response to connectivity provided by corridors, whereas species richness of wind-dispersed plants was unaffected by connectivity and increased in response to changes in patch shape--higher edge-to-interior ratio--created by corridors. Corridors promoted biodiversity spillover for native species of the threatened longleaf pine ecosystem being restored in our experiment, but not for exotic species. By extending economically driven spillover concepts from marine fisheries and crop pollination systems, we show how reconnecting landscapes amplifies biodiversity conservation both within and beyond reserve borders.

A conceptual model of movement ecology has recently been advanced to explain all movement by considering the interaction of four elements: internal state, motion capacity, navigation capacities, and external factors. We modified this framework to generate predictions for species richness dynamics of fragmented plant communities and tested them in experimental landscapes across a 7-year time series. We found that two external factors, dispersal vectors and habitat features, affected species colonization and recolonization in habitat fragments and their effects varied and depended on motion capacity. Bird-dispersed species richness showed connectivity effects that reached an asymptote over time, but no edge effects, whereas wind-dispersed species richness showed steadily accumulating edge and connectivity effects, with no indication of an asymptote. Unassisted species also showed increasing differences caused by connectivity over time, whereas edges had no effect. Our limited use of proxies for movement ecology (e.g., dispersal mode as a proxy for motion capacity) resulted in moderate predictive power for communities and, in some cases, highlighted the importance of a more complete understanding of movement ecology for predicting how landscape conservation actions affect plant community dynamics.