Harnessing ecological theory to enhance ecosystem restoration.

Ecosystem restoration can increase the health and resilience of nature and humanity. As a result, the international community is championing habitat restoration as a primary solution to address the dual climate and biodiversity crises. Yet most ecosystem restoration efforts to date have underperformed, failed, or been burdened by high costs that prevent upscaling. To become a primary, scalable conservation strategy, restoration efficiency and success must increase dramatically. Here, we outline how integrating ten foundational ecological theories that have not previously received much attention - from hierarchical facilitation to macroecology - into ecosystem restoration planning and management can markedly enhance restoration success. We propose a simple, systematic approach to determining which theories best align with restoration goals and are most likely to bolster their success. Armed with a century of advances in ecological theory, restoration practitioners will be better positioned to more cost-efficiently and effectively rebuild the world's ecosystems and support the resilience of our natural resources.

Species eradication for ecosystem restoration.

Many of the world's ecosystems are under unprecedented stress as human pressures have escalated to be a dominant driver of ecosystem composition and condition. Direct impacts such as agriculture, extraction, and development are impacting vast swathes of land and ocean, while the effects of human-caused climate change are felt even in the most remote parts of marine and terrestrial wildernesses. These impacts are resulting in changes ranging from ecosystem collapse or replacement to novel mixes of species due to temperature-driven range shifts. While reducing human pressures is paramount for the future viability of vulnerable ecosystems, much attention is now also focused on whether degraded areas can be restored. Indeed, the UN has declared 2021-2030 the Decade on Ecosystem Restoration, which aims to "prevent, halt and reverse the degradation of ecosystems on every continent and in every ocean".

Functional consequences of animal community changes in managed grasslands: An application of the CAFE approach.

In the midst of an ongoing biodiversity crisis, much research has focused on species losses and their impacts on ecosystem functioning. The functional consequences (ecosystem response) of shifts in communities are shaped not only by changes in species richness, but also by compositional shifts that result from species losses and gains. Species differ in their contribution to ecosystem functioning, so species identity underlies the consequences of species losses and gains on ecosystem functions. Such research is critical to better predict the impact of disturbances on communities and ecosystems. We used the "Community Assembly and the Functioning of Ecosystems" (CAFE) approach, a modification of the Price equation to understand the functional consequences and relative effects of richness and composition changes in small nonvolant mammal and dung beetle communities as a result of two common disturbances in North American prairie restorations, prescribed fire and the reintroduction of large grazing mammals. Previous research in this system has shown dung beetles are critically important decomposers, while small mammals modulate much energy in prairie food webs. We found that dung beetle communities were more responsive to bison reintroduction and prescribed fires than small nonvolant mammals. Dung beetle richness increased after bison reintroduction, with higher dung beetle community biomass resulting from changes in remaining species (context-dependent component) rather than species turnover (richness components); prescribed fire caused a minor increase in dung beetle biomass for the same reason. For small mammals, bison reintroduction reduced energy transfer through the loss of species, while prescribed fire had little impact on either small mammal richness or energy transfer. The CAFE approach demonstrates how bison reintroduction controls small nonvolant mammal communities by increasing prairie food web complexity, and increases dung beetle populations with possible benefits for soil health through dung mineralization and soil bioturbation. Prescribed fires, however, have little effect on small mammals and dung beetles, suggesting a resilience to fire. These findings illustrate the key role of re-establishing historical disturbance regimes when restoring endangered prairie ecosystems and their ecological function.

Herbivory limits success of vegetation restoration globally.

Restoring vegetation in degraded ecosystems is an increasingly common practice for promoting biodiversity and ecological function, but successful implementation is hampered by an incomplete understanding of the processes that limit restoration success. By synthesizing terrestrial and aquatic studies globally (2594 experimental tests from 610 articles), we reveal substantial herbivore control of vegetation under restoration. Herbivores at restoration sites reduced vegetation abundance more strongly (by 89%, on average) than those at relatively undegraded sites and suppressed, rather than fostered, plant diversity. These effects were particularly pronounced in regions with higher temperatures and lower precipitation. Excluding targeted herbivores temporarily or introducing their predators improved restoration by magnitudes similar to or greater than those achieved by managing plant competition or facilitation. Thus, managing herbivory is a promising strategy for enhancing vegetation restoration efforts.

An invasive appetite: Combining molecular and stable isotope analyses to reveal the diet of introduced house mice (Mus musculus) on a small, subtropical island.

House mice (Mus musculus) pose a conservation threat on islands, where they adversely affect native species' distributions, densities, and persistence. On Sand Island of Kuaihelani, mice recently began to depredate nesting adult moli (Laysan Albatross, Phoebastria immutabilis). Efforts are underway to eradicate mice from Sand Island, but knowledge of mouse diet is needed to predict ecosystem response and recovery following mouse removal. We used next-generation sequencing to identify what mice eat on Sand Island, followed by stable isotope analysis to estimate the proportions contributed by taxa to mouse diet. We collected paired fecal and hair samples from 318 mice between April 2018 to May 2019; mice were trapped approximately every eight weeks among four distinct habitat types to provide insight into temporal and spatial variation. Sand Island's mice mainly consume arthropods, with nearly equal (but substantially smaller) contributions of C3 plants, C4 plants, and moli. Although seabird tissue is a small portion of mouse diet, mice consume many detrital-feeding arthropods in and around seabird carcasses, such as isopods, flesh flies, ants, and cockroaches. Additionally, most arthropods and plants eaten by mice are non-native. Mouse diet composition differs among habitat types but changes minimally throughout the year, indicating that mice are not necessarily limited by food source availability or accessibility. Eradication of house mice may benefit seabirds on Sand Island (by removing a terrestrial, non-native predator), but it is unclear how arthropod and plant communities may respond and change. Non-native and invasive arthropods and plants previously consumed (and possibly suppressed) by mice may be released post-eradication, which could prevent recovery of native taxa. Comprehensive knowledge of target species' diet is a critical component of eradication planning. Dietary information should be used both to identify and to monitor which taxa may respond most strongly to invasive species removal and to assess if proactive, pre-eradication management activities are warranted.

Hold big business to task on ecosystem restoration.

Corporate reporting must embrace holistic, scientific principles.

Tracking the global application of conservation translocation and social attraction to reverse seabird declines.

The global loss of biodiversity has inspired actions to restore nature across the planet. Translocation and social attraction actions deliberately move or lure a target species to a restoration site to reintroduce or augment populations and enhance biodiversity and ecosystem resilience. Given limited conservation funding and rapidly accelerating extinction trajectories, tracking progress of these interventions can inform best practices and advance management outcomes. Seabirds are globally threatened and commonly targeted for translocation and social attraction ("active seabird restoration"), yet no framework exists for tracking these efforts nor informing best practices. This study addresses this gap for conservation decision makers responsible for seabirds and coastal management. We systematically reviewed active seabird restoration projects worldwide and collated results into a publicly accessible Seabird Restoration Database. We describe global restoration trends, apply a systematic process to measure success rates and response times since implementation, and examine global factors influencing outcomes. The database contains 851 active restoration events in 551 locations targeting 138 seabird species; 16% of events targeted globally threatened taxa. Visitation occurred in 80% of events and breeding occurred in 76%, on average 2 y after implementation began (SD = 3.2 y). Outcomes varied by taxonomy, with the highest and quickest breeding response rates for Charadriiformes (terns, gulls, and auks), primarily with social attraction. Given delayed and variable response times to active restoration, 5 y is appropriate before evaluating outcomes. The database and results serve as a model for tracking and evaluating restoration outcomes, and is applicable to measuring conservation interventions for additional threatened taxa.

Reintroduced megaherbivores indirectly shape small-mammal responses to moonlight.

Moonlight structures activity patterns of many nocturnal species. Bright moonlight often limits the activity of nocturnal prey, but dense vegetation weakens this effect. Using 8 years of live-trapping data, we asked whether reintroduced megaherbivores (Bison bison) indirectly altered moonlight avoidance by small mammals in tallgrass prairies. In plots with bison, plants intercepted 20% less light, allowing more moonlight to reach ground level. During nights with no moonlight, Peromyscus maniculatus activity was similar in plots with and without bison. During nights with peak moonlight, P. maniculatus activity was four times greater in plots without bison compared to plots with bison. Conversely, Microtus ochrogaster activity was twice as great during full moons compared to new moons, but only in plots with bison. We also equipped a subset of traps with temperature sensors to estimate trap-entry time. Although M. ochrogaster was more active on bright nights, most activity occurred before moonrise or after moonset, avoiding periods of bright moonlight. We conclude that megaherbivores play an unappreciated but important indirect role in tallgrass prairies by inducing behavioral shifts in other animal species. Because overlap in activity patterns can predict the likelihood of predator-prey encounters, such activity shifts have important implications for trophic interactions throughout restored prairie food webs. Additional work to understand interspecific and intraspecific variation in response to moonlight may improve efforts to forecast changes in community assembly due to restoration and land-use change.

The effect of seabird presence and seasonality on ground-active spider communities across temperate islands.

Seabirds influence island ecosystems through nutrient additions and physical disturbance. These influences can have opposing effects on an island's invertebrate predator populations. Spiders (order: Araneae) are an important predator in many terrestrial island ecosystems, yet little is known about how seabird presence influences spider communities at the intraisland scale, or how they respond to seasonality in seabird colony attendance.We investigated the effects of seabird presence and seasonality on ground-active spider community structure (activity-density, family-level richness, age class, and sex structure) and composition at the family-level across five short-tailed shearwater breeding islands around south-eastern Tasmania, Australia. Using 75 pitfall traps (15 per island), spiders were collected inside, near, and outside seabird colonies on each island, at five different stages of the short-tailed shearwater breeding cycle over a year. Pitfall traps were deployed for a total of 2674 days, capturing 1592 spiders from 26 families with Linyphiidae and Lycosidae the most common. Spider activity-density was generally greater inside than outside seabird colonies, while family-level richness was generally higher outside seabird colonies. For these islands, seabird breeding stage did not affect activity-densities, but there were some seasonal changes in age class and sex structures with more adult males captured during winter. Our results provide some of the first insights into the spatial and temporal influences seabirds have on spider communities. We also provide some of the first records of spider family occurrences for south-eastern Tasmanian islands, which will provide an important baseline for assessing future change.

Harnessing island-ocean connections to maximize marine benefits of island conservation.

Islands support unique plants, animals, and human societies found nowhere else on the Earth. Local and global stressors threaten the persistence of island ecosystems, with invasive species being among the most damaging, yet solvable, stressors. While the threat of invasive terrestrial mammals on island flora and fauna is well recognized, recent studies have begun to illustrate their extended and destructive impacts on adjacent marine environments. Eradication of invasive mammals and restoration of native biota are promising tools to address both island and ocean management goals. The magnitude of the marine benefits of island restoration, however, is unlikely to be consistent across the globe. We propose a list of six environmental characteristics most likely to affect the strength of land-sea linkages: precipitation, elevation, vegetation cover, soil hydrology, oceanographic productivity, and wave energy. Global databases allow for the calculation of comparable metrics describing each environmental character across islands. Such metrics can be used today to evaluate relative potential for coupled land-sea conservation efforts and, with sustained investment in monitoring on land and sea, can be used in the future to refine science-based planning tools for integrated land-sea management. As conservation practitioners work to address the effects of climate change, ocean stressors, and biodiversity crises, it is essential that we maximize returns from our management investments. Linking efforts on land, including eradication of island invasive mammals, with marine restoration and protection should offer multiplied benefits to achieve concurrent global conservation goals.

Arthropod Predation of Vertebrates Structures Trophic Dynamics in Island Ecosystems.

AbstractOn isolated islands, large arthropods can play an important functional role in ecosystem dynamics. On the Norfolk Islands group, South Pacific, we monitored the diet and foraging activity of an endemic chilopod, the Phillip Island centipede (Cormocephalus coynei), and used a stable isotope mixing model to estimate dietary proportions. Phillip Island centipede diet is represented by vertebrate animals (48%) and invertebrates (52%), with 30.5% consisting of squamates, including the Lord Howe Island skink (Oligosoma lichenigera) and Günther's island gecko (Christinus guentheri); 7.9% consisting of black-winged petrel (Pterodroma nigripennis) nestlings; and 9.6% consisting of marine fishes scavenged from regurgitated seabird meals. Centipede predation was the principal source of petrel nestling mortality, with annual rates of predation varying between 11.1% and 19.6% of nestlings. This means that 2,109-3,724 black-winged petrel nestlings may be predated by centipedes annually. Petrels produce a single offspring per year; therefore, predation of nestlings by centipedes represents total breeding failure for a pair in a given year. Our work demonstrates that arthropods can play a leading role in influencing vertebrate reproductive output and modifying trophic structures and nutrient flow in island ecosystems.

Disturbance-Induced Trophic Niche Shifts In Ground Beetles (Coleoptera: Carabidae) In Restored Grasslands.

Ecosystem restoration is a critical component of land management, countering the loss of native biodiversity. Restoration efforts are enhanced by reintroducing naturally occurring ecosystem processes, including disturbances that may impact species characteristics such as niche position or niche size. In grasslands, grazing and fire affect plant diversity and habitat complexity, which potentially influence insect dietary behaviors and thus their contributions to functions like seed and arthropod predation. Using carbon and nitrogen stable isotopes, we characterized variation in the dietary niche of six ground beetle species (Coleoptera: Carabidae) in response to grazing by reintroduced bison and prescribed fire disturbances in twenty tallgrass prairies. Management disturbances did not affect activity density for most beetle species and mean trophic position was mostly unaffected. However, five of six species exhibited increased trophic niche area and breadth with disturbances, indicating a switch to a more generalist diet that incorporated a wider range of food items. The combination of bison and fire impacts may increase vegetation patchiness and heterogeneity, driving these diet changes. Morphological traits and microhabitat preferences might mediate response to disturbances and the resulting heterogeneity. Combining prescribed fire and grazing, which increases plant diversity and vegetation structural diversity, may help beetle communities establish over time and support the ecological functions to which these insects contribute.

Effects of management outweigh effects of plant diversity on restored animal communities in tallgrass prairies.

A primary goal of ecological restoration is to increase biodiversity in degraded ecosystems. However, the success of restoration ecology is often assessed by measuring the response of a single functional group or trophic level to restoration, without considering how restoration affects multitrophic interactions that shape biodiversity. An ecosystem-wide approach to restoration is therefore necessary to understand whether animal responses to restoration, such as changes in biodiversity, are facilitated by changes in plant communities (plant-driven effects) or disturbance and succession resulting from restoration activities (management-driven effects). Furthermore, most restoration ecology studies focus on how restoration alters taxonomic diversity, while less attention is paid to the response of functional and phylogenetic diversity in restored ecosystems. Here, we compared the strength of plant-driven and management-driven effects of restoration on four animal communities (ground beetles, dung beetles, snakes, and small mammals) in a chronosequence of restored tallgrass prairie, where sites varied in management history (prescribed fire and bison reintroduction). Our analyses indicate that management-driven effects on animal communities were six-times stronger than effects mediated through changes in plant biodiversity. Additionally, we demonstrate that restoration can simultaneously have positive and negative effects on biodiversity through different pathways, which may help reconcile variation in restoration outcomes. Furthermore, animal taxonomic and phylogenetic diversity responded differently to restoration, suggesting that restoration plans might benefit from considering multiple dimensions of animal biodiversity. We conclude that metrics of plant diversity alone may not be adequate to assess the success of restoration in reassembling functional ecosystems.

Reintroduced grazers and prescribed fire effects on beetle assemblage structure and function in restored grasslands.

Ecological restoration seeks to reestablish functioning ecosystems, but planning and evaluation often focus on taxonomic community structure and neglect consumers and their functional roles. The functional trait composition of insect assemblages, which make up the majority of animal diversity in many systems, can reveal how they are affected by restoration management and the consequences for ecosystem function. We sampled ground beetle (Coleoptera: Carabidae) assemblages in restored tallgrass prairies varying in management with prescribed fire and reintroduced American bison (Bison bison) to describe their taxonomic and functional trait structure. We also measured seed and arthropod predation to relate management, beetle assemblage characteristics, and function, and to test if function is maximized by trait diversity, dominant trait values, or beetle abundance. Beetle assemblages primarily varied with restoration age, declining over time in richness and both taxonomic and functional diversity, but bison presence also influenced taxonomic composition. Prescribed fire reduced seed predation in summer and arthropod predation in fall. Although seed predation was unrelated to beetle assemblages, arthropod predation was greater in sites with higher abundances of carnivorous ground beetles. The relatively weak impacts of fire and bison on functional assemblage structure is a promising sign that these management disturbances, aimed at supporting a diverse native plant community, are not detrimental to beetle assemblages. The significance of reduced predator function following prescribed fire will depend on the restoration context and whether seed or arthropod predation relates to management goals.

Global hotspots for coastal ecosystem-based adaptation.

Helping the world's coastal communities adapt to climate change impacts requires evaluating the vulnerability of coastal communities and assessing adaptation options. This includes understanding the potential for 'natural' infrastructure (ecosystems and the biodiversity that underpins them) to reduce communities' vulnerability, alongside more traditional 'hard' infrastructure approaches. Here we present a spatially explicit global evaluation of the vulnerability of coastal-dwelling human populations to key climate change exposures and explore the potential for coastal ecosystems to help people adapt to climate change (ecosystem-based adaptation (EbA)). We find that mangroves and coral reefs are particularly well situated to help people cope with current weather extremes, a function that will only increase in importance as people adapt to climate change now and in coming decades. We find that around 30.9 million people living within 2km of the coast are highly vulnerable to tropical storms and sea-level rise (SLR). Mangroves and coral reefs overlap these threats to at least 5.3 and 3.4 million people, respectively, with substantial potential to dissipate storm surges and improve resilience against SLR effects. Significant co-benefits from mangroves also accrue, with 896 million metric tons of carbon stored in their soils and above- and below-ground biomass. Our framework offers a tool for prioritizing 'hotspots' of coastal EbA potential for further, national and local analyses to quantify risk reduction and, thereby, guide investment in coastal ecosystems to help people adapt to climate change. In doing so, it underscores the global role that conserving and restoring ecosystems can play in protecting human lives and livelihoods, as well as biodiversity, in the face of climate change.

The geography of biodiversity change in marine and terrestrial assemblages.

Human activities are fundamentally altering biodiversity. Projections of declines at the global scale are contrasted by highly variable trends at local scales, suggesting that biodiversity change may be spatially structured. Here, we examined spatial variation in species richness and composition change using more than 50,000 biodiversity time series from 239 studies and found clear geographic variation in biodiversity change. Rapid compositional change is prevalent, with marine biomes exceeding and terrestrial biomes trailing the overall trend. Assemblage richness is not changing on average, although locations exhibiting increasing and decreasing trends of up to about 20% per year were found in some marine studies. At local scales, widespread compositional reorganization is most often decoupled from richness change, and biodiversity change is strongest and most variable in the oceans.

Restoration and repair of Earth's damaged ecosystems.

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.

A global review of past land use, climate, and active vs. passive restoration effects on forest recovery.

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.