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What is a biocrust? A refined, contemporary definition for a broadening research community

Studies of biological soil crusts (biocrusts) have proliferated over the last few decades. The biocrust literature has broadened, with more studies assessing and describing the function of a variety of biocrust communities in a broad range of biomes and habitats and across a large spectrum of disciplines, and also by the incorporation of biocrusts into global perspectives and biogeochemical models. As the number of biocrust researchers increases, along with the scope of soil communities defined as ‘biocrust’, it is worth asking whether we all share a clear, universal, and fully articulated definition of what constitutes a biocrust. In this review, we synthesize the literature with the views of new and experienced biocrust researchers, to provide a refined and fully elaborated definition of biocrusts. In doing so, we illustrate the ecological relevance and ecosystem services provided by them. We demonstrate that biocrusts are defined by four distinct elements: physical structure, functional characteristics, habitat, and taxonomic composition. We describe outgroups, which have some, but not all, of the characteristics necessary to be fully consistent with our definition and thus would not be considered biocrusts. We also summarize the wide variety of different types of communities that fall under our definition of biocrusts, in the process of highlighting their global distribution. Finally, we suggest the universal use of the Belnap, Büdel & Lange definition, with minor modifications: Biological soil crusts (biocrusts) result from an intimate association between soil particles and differing proportions of photoautotrophic (e.g. cyanobacteria , algae , lichens , bryophytes) and heterotrophic (e.g. bacteria , fungi , archaea) organisms , which live within , or immediately on top of , the uppermost millimetres of soil. Soil particles are aggregated through the presence and activity of these often extremotolerant biota that desiccate regularly , and the resultant living crust covers the surface of the ground as a coherent layer. With this detailed definition of biocrusts, illustrating their ecological functions and widespread distribution, we hope to stimulate interest in biocrust research and inform various stakeholders (e.g. land managers, land users) on their overall importance to ecosystem and Earth system functioning.

Biological Reviews

Projected climate and land use changes drive plant community composition in agricultural wetlands

Playa wetlands in the Great Plains, USA support a wide variety of plant species not found elsewhere in this agriculturally-dominated region due to the ephemeral presence of standing water and hydric soils within playas. If longer dry periods occur due to climate change or if changes in surrounding land use alter sediment accumulation rates and water storage capacity in playas, plant communities could experience decreased diversity, with lasting effects on ecosystem services provided by playas in the Great Plains and at a continental-level in North America. We quantified potential changes in playa wetland plant community composition associated with predicted changes in precipitation and land use in the Great Plains through the end of the 21 st century. We conducted two six-month greenhouse experiments mimicking field conditions using intact mesocosms collected from playas in Nebraska and Texas. In the precipitation experiment, treatments derived from historical precipitation observations and three future moderate emissions (CMIP5 RCP4.5) downscaled climate projections were applied to mesocosms. For the land use experiment, treatments were simulated by nitrogen (N) applications to soil ranging from 0 to 100 mg-N L -1 with each precipitation event under historical rainfall patterns, representing increasing and decreasing area in agricultural use in playa watersheds. Plant communities tended to shift toward more native species under projected future climate conditions, but as N runoff increased, native species richness decreased. Agricultural land-use surrounding playas may have a greater effect on wetland plant communities than future alterations to hydrology based on climate change in the Great Plains; thus, efforts to reduce nutrient runoff into playas would likely mitigate loss in ecosystem function in the coming decades.

Nebraska, Texas

Fire and tree death: Understanding and improving modeling of fire-induced tree mortality

Each year wildland fires kill and injure trees on millions of forested hectares globally, affecting plant and animal biodiversity, carbon storage, hydrologic processes, and ecosystem services. The underlying mechanisms of fire-caused tree mortality remain poorly understood, however, limiting the ability to accurately predict mortality and develop robust modeling applications, especially under novel future climates. Virtually all post-fire tree mortality prediction systems are based on the same underlying empirical model described in Ryan and Reinhardt (1988 Can. J. For. Res. 18 1291–7), which was developed from a limited number of species, stretching model assumptions beyond intended limits. We review the current understanding of the mechanisms of fire-induced tree mortality, provide recommended standardized terminology, describe model applications and limitations, and conclude with key knowledge gaps and future directions for research. We suggest a two-pronged approach to future research: (1) continued improvements and evaluations of empirical models to quantify uncertainty and incorporate new regions and species and (2) acceleration of basic, physiological research on the proximate and ultimate causes of fire-induced tree mortality to incorporate processes of tree death into models. Advances in both empirical and process fire-induced tree modeling will allow creation of hybrid models that could advance understanding of how fire injures and kills trees, while improving prediction accuracy of fire-driven feedbacks on ecosystems and landscapes, particularly under novel future conditions.

Environmental Research Letters

On the sustainability of inland fisheries: Finding a future for the forgotten

At present, inland fisheries are not often a national or regional governance priority and as a result, inland capture fisheries are undervalued and largely overlooked. As such they are threatened in both developing and developed countries. Indeed, due to lack of reliable data, inland fisheries have never been part of any high profile global fisheries assessment and are notably absent from the Sustainable Development Goals. The general public and policy makers are largely ignorant of the plight of freshwater ecosystems and the fish they support, as well as the ecosystem services generated by inland fisheries. This ignorance is particularly salient given that the current emphasis on the food-water-energy nexus often fails to include the important role that inland fish and fisheries play in food security and supporting livelihoods in low-income food deficit countries. Developing countries in Africa and Asia produce about 11 million tonnes of inland fish annually, 90 % of the global total. The role of inland fisheries goes beyond just kilocalories; fish provide important micronutrients and essentially fatty acids. In some regions, inland recreational fisheries are important, generating much wealth and supporting livelihoods. The following three key recommendations are necessary for action if inland fisheries are to become a part of the food-water-energy discussion: invest in improved valuation and assessment methods, build better methods to effectively govern inland fisheries (requires capacity building and incentives), and develop approaches to managing waters across sectors and scales. Moreover, if inland fisheries are recognized as important to food security, livelihoods, and human well-being, they can be more easily incorporated in regional, national, and global policies and agreements on water issues. Through these approaches, inland fisheries can be better evaluated and be more fully recognized in broader water resource and aquatic ecosystem planning and decision-making frameworks, enhancing their value and sustainability for the future.

Ambio

Herbaceous production lost to tree encroachment in United States rangelands

Rangelands of the United States provide ecosystem services that benefit society and rural economies. Native tree encroachment is often overlooked as a primary threat to rangelands due to the slow pace of tree cover expansion and the positive public perception of trees. Still, tree encroachment fragments these landscapes and reduces herbaceous production, thereby threatening habitat quality for grassland wildlife and the economic sustainability of animal agriculture. Recent innovations in satellite remote sensing permit the tracking of tree encroachment and the corresponding impact on herbaceous production. We analysed tree cover change and herbaceous production across the western United States from 1990 to 2019. We show that tree encroachment is widespread in US rangelands; absolute tree cover has increased by 50% (77,323 km 2 ) over 30 years, with more than 25% (684,852 km 2 ) of US rangeland area experiencing tree cover expansion. Since 1990, 302 ± 30 Tg of herbaceous biomass have been lost. Accounting for variability in livestock biomass utilization and forage value reveals that this lost production is valued at between $4.1–$5.6 billion US dollars. Synthesis and applications . The magnitude of impact of tree encroachment on rangeland loss is similar to conversion to cropland, another well-known and primary mechanism of rangeland loss in the US Prioritizing conservation efforts to prevent tree encroachment can bolster ecosystem and economic sustainability, particularly among privately-owned lands threatened by land-use conversion.

Journal of Applied Ecology

Measuring carbon and nitrogen bioassimilation, burial, and denitrification contributions of oyster reefs in Gulf coast estuaries

The eastern oyster ( Crassostrea virginica ) and the reefs they create provide significant ecosystem services. This study measured their possible role in nutrient mitigation through bioassimilation, burial, and oyster-mediated sediment denitrification in near-shore shallow water (< 1 m water depth) and deep-water (> 1 m water depth) oyster reefs in Louisiana. Nitrogen (N) and carbon (C) in shell and tissue differed by oyster reproductive status, size, and habitat type. Changes in tissue percent N and C post-spawning combined with significant reductions in tissue dry weight from the release of gametes, resulted in 20 and 46% reductions in tissue N and C load (mg), respectively, for a 100-mm oyster. Oyster reefs did not enhance burial rates, with burial range rates estimated at 1.4–2.6 g N m −2 year −1 , and 26.9–43.8 g C m −2 year −1 . Closed-system ex situ incubations indicated net denitrification in all habitat types studied, with the highest rates exceeding 600 µmol N m −2 h −1 during the summer, but no enhancement attributable to oyster reefs specifically. Within the highly productive, organic-rich wetland complex systems of coastal Louisiana, oyster reefs were not associated with enhanced denitrification, likely due to the organic-rich setting, and redundant supplies of organic nitrogen and carbon from adjacent marshes. Context remains critical in determining ecosystem provision of habitats, and efforts to extrapolate and predict nitrogen removal across locations necessitates consideration of local conditions. Considering the large extent of reefs and oyster production across coastal Louisiana, oyster habitats may still contribute to N and C mitigation, but their unique contribution likely comes from bioassimilation, and removal of the oysters from the system.

Louisiana

Soil carbon consequences of historic hydrologic impairment and recent restoration in coastal wetlands

Coastal wetlands provide key ecosystem services, including substantial long-term storage of atmospheric CO2 in soil organic carbon pools. This accumulation of soil organic matter is a vital component of elevation gain in coastal wetlands responding to sea-level rise. Anthropogenic activities that alter coastal wetland function through disruption of tidal exchange and wetland water levels are ubiquitous. This study assesses soil vertical accretion and organic carbon accretion across five coastal wetlands that experienced over a century of impounded hydrology, followed by restoration of tidal exchange 5 to 14 years prior to sampling. Nearby marshes that never experienced tidal impoundment served as controls with natural hydrology to assess the impact of impoundment and restoration. Dated soil cores indicate that elevation gain and carbon storage were suppressed 30–70 % during impoundment, accounting for the majority of elevation deficit between impacted and natural sites. Only one site had substantial subsidence, likely due to oxidation of soil organic matter. Vertical and carbon accretion gains were achieved at all restored sites, with carbon burial increasing from 96 ± 33 to 197 ± 64 g C m −2 y −1 . The site with subsidence was able to accrete at double the rate (13 ± 5.6 mm y −1 ) of the natural complement, due predominantly to organic matter accumulation rather than mineral deposition, indicating these ecosystems are capable of large dynamic responses to restoration when conditions are optimized for vegetation growth. Hydrologic restoration enhanced elevation resilience and climate benefits of these coastal wetlands.

Massachusetts

A classification of streamflow patterns across the coastal Gulf of Alaska

Streamflow controls many freshwater and marine processes, including salinity profiles, sediment composition, fluxes of nutrients, and the timing of animal migrations. Watersheds that border the Gulf of Alaska (GOA) comprise over 400,000 km 2 of largely pristine freshwater habitats and provide ecosystem services such as reliable fisheries for local and global food production. Yet no comprehensive watershed-scale description of current temporal and spatial patterns of streamflow exists within the coastal GOA. This is an immediate need because the spatial distribution of future streamflow patterns may shift dramatically due to warming air temperature, increased rainfall, diminishing snowpack, and rapid glacial recession. Our primary goal was to describe variation in streamflow patterns across the coastal GOA using an objective set of descriptors derived from flow predictions at the downstream-most point within each watershed. We leveraged an existing hydrologic runoff model and Bayesian mixture model to classify 4,140 watersheds into 13 classes based on seven streamflow statistics. Maximum discharge timing (annual phase shift) and magnitude relative to mean discharge (amplitude) were the most influential attributes. Seventy-six percent of watersheds by number showed patterns consistent with rain or snow as dominant runoff sources, while the remaining watersheds were driven by rain-snow, glacier, or low-elevation wetland runoff. Streamflow classes exhibited clear mechanistic links to elevation, ice coverage, and other landscape features. Our classification identifies watersheds that might shift streamflow patterns in the near future and, importantly, will help guide the design of studies that evaluate how hydrologic change will influence coastal GOA ecosystems.

Alaska

Predictive regressive models of recent marsh sediment thickness improve the quantification of coastal marsh sediment budgets

Coastal marsh wetlands experience variations in vertical gains and losses through time, which have allowed them to infill relict topography and record variations in drivers. The stratigraphic unit associated with the development of the marsh also reflects the long-term importance of key ecosystem services supplied by the marsh environment, including carbon storage and storm mitigation. Mapping these coastal wetland sediments and the marsh unit thickness is challenging as traditional coastal geophysical tools are not easily deployable (acoustic methods) or are unreliable in saline-soil environments (e.g., ground-penetrating radar), leaving core-based methods the most viable mapping method. In the present study, we utilized prior information on the geologic architecture of the region to select spatial and physical metrics that likely persisted throughout evolution of the marsh during the late Holocene. We then assessed the individual and collective power of these metrics to predict marsh thickness observed from cores. Employing regressive predictive models powered by these data, we improve the quantification of marsh thickness for a coastal fringing marsh within the Grand Bay estuary in Mississippi and Alabama (USA). The information gained from this approach yields improved estimates of the carbon stocks in this environment. Additionally, the stored sediment masses reflect the past, and potential future, persistence of the Grand Bay marsh under historical and present marsh-estuarine sediment exchange fluxes. Such improvements to both the sediment budget of recent marsh stratigraphic units and the spatial extent provide new resources for comparison with large-scale landscape models, the latter of which may be used, when validated, to predict future change and ecosystem transformations.

Alabama. Mississippi

Confluence of time and space: An innovation for quantifying dynamics of hydrologic floodplain connectivity with remote sensing and GIS

Hydrologic connectivity is a crucial determinant of aquatic ecosystem services, governing the exchange of nutrients, sediments, chemicals, and biota. Various indices and metrics exist for quantifying hydrologic connectivity across diverse environments and scales. However, existing methodologies often fail to adequately capture lateral connectivity between floodplain lakes and streams across vast, low-relief, multi-lake floodplain systems. This study introduces a novel approach for quantifying lateral hydrologic connectivity specifically tailored for floodplain lakes connecting to streams within the expansive floodplain of the Lower Mississippi River. This approach centers on the spatial and temporal intersection of lakes and streams, leveraging remote sensing and GIS data to estimate nine distinct metrics of hydrologic connectivity. To assess the reliability of the method, the study estimated connectivity metrics for 92 randomly selected floodplain lakes, comprising 53 lakes connected to large streams (Strahler order >7), 13 lakes connected to medium (order 4-6) streams, and 26 lakes connected to small (order 1-3) streams. As expected, there was significant variability in hydrologic connectivity across different stream size classes. The outlined approach contributes valuable insights into the hydrologic connectivity of floodplain lakes and offers a generalizable framework applicable to other floodplains. Its versatility makes it a practical tool for understanding connectivity requirements for biota and facilitating applications in conservation and water resources management. Thus, this work represents a meaningful step toward advancing our understanding of lateral hydrologic connectivity dynamics in complex aquatic ecosystems.

Arkansas, Illinois, Kentucky, Louisiana, Mississip

Knowledge sharing for shared success in the decade on ecosystem restoration

The Decade on Ecosystem Restoration aims to provide the means and incentives for upscaling restoration efforts worldwide. Although ecosystem restoration is a broad, interdisciplinary concept, effective ecological restoration requires sound ecological knowledge to successfully restore biodiversity and ecosystem services in degraded landscapes. We emphasize the critical role of knowledge and data sharing to inform synthesis for the most robust restoration science possible. Such synthesis is critical for helping restoration ecologists better understand how context affects restoration outcomes, and to increase predictive capacity of restoration actions. This predictive capacity can help to provide better information for evidence-based decision-making, and scale-up approaches to meet ambitious targets for restoration. We advocate for a concerted effort to collate species-level, fine-scale, ecological community data from restoration studies across a wide range of environmental and ecological gradients. Well-articulated associated metadata relevant to experience and social or landscape contexts can further be used to explain outcomes. These data could be carefully curated and made openly available to the restoration community to help to maximize evidence-based knowledge sharing, enable flexible re-use of existing data and support predictive capacity in ecological community responses to restoration actions. We detail how integrated data, analysis and knowledge sharing via synthesis can support shared success in restoration ecology by identifying successful and unsuccessful outcomes across diverse systems and scales. We also discuss potential interdisciplinary solutions and approaches to overcome challenges associated with bringing together subfields of restoration practice. Sharing this knowledge and data openly can directly inform actions and help to improve outcomes for the Decade on Ecosystem Restoration.

Ecological Solutions and Evidence

Evaluation of stream and wetlands restoration using UAS-based thermal infrared mapping

Large-scale wetland restoration often focuses on repairing the hydrologic connections degraded by anthropogenic modifications. Of these hydrologic connections, groundwater discharge is an important target, as these surface water ecosystem control points are important to thermal stability, among other ecosystem services. However, evaluating the effectiveness of the restoration activities on establishing groundwater discharge connection is often difficult over the vast area and often challenging or inaccessible terrain of wetlands. Unoccupied aerial systems (UAS) are now routinely used for collecting aerial imagery and creating digital surface models (DSM). Lightweight thermal infrared (TIR) sensors provide another payload option for generation of sub-meter resolution aerial TIR orthophotos. This technology allows for the rapid and safe survey of groundwater discharge areas. Aerial TIR water-surface data were collected March 2019 at Tidmarsh Farms, a former commercial cranberry peatland located in coastal Massachusetts, USA (41°54'17.6"N 70°34'17.4"W), where stream and wetland restoration actions were completed in 2016. Here we present a 0.4 km2 georeferenced, temperature calibrated TIR orthophoto of the area. The image represents a mosaic of nearly 900 TIR images captured by UAS in a single morning with a total flight time of 36 minutes, and is supported by a DSM derived from UAS visible imagery. The survey was conducted in winter to maximize temperature contrast between relatively warm groundwater and colder ambient surface environment; lower-density groundwater rises above cool surface waters and thus can be imaged by a UAS. The resulting TIR orthomosaic shows fine detail of seepage distribution and downstream influence along the several restored channel forms, which was an objective of the ecological restoration design. The restored stream channel has increased connectivity to peatland groundwater discharge, reducing the ecosystem thermal stressors. Such aerial techniques can be used to guide ecological restoration design and assess post-restoration outcomes, especially in settings where ecosystem structure and function is governed by groundwater and surface water interaction.

Water

A biological condition gradient for coral reefs in the US Caribbean Territories: Part I. Coral narrative rules

As coral reef condition and sustainability continue to decline worldwide, losses of critical habitat and their ecosystem services have generated an urgency to understand and communicate reef response to management actions, environmental contamination, and natural disasters. Increasingly, coral reef protection and restoration programs emphasize the need for robust assessment tools for protecting high-quality waters and establishing conservation goals. Of equal importance is the need to communicate assessment results to stakeholders, beneficiaries, and the public so that environmental consequences of decisions are understood. The Biological Condition (BCG) model provides a structure to evaluate the condition of a coral reef in increments of change along a gradient of human disturbance. Communication of incremental change, regardless of direction, is important for decision makers and the public to better understand what is gained or lost depending on what actions are taken. We developed a narrative (qualitative) Biological Condition Gradient (BCG) from the consensus of a diverse expert panel to provide a framework for coral reefs in US Caribbean Territories. The model uses narrative descriptions of biological attributes for benthic organisms to evaluate reefs relative to undisturbed or minimally disturbed conditions. Using expert elicitation, narrative decision rules were proposed and deliberated to discriminate among six levels of change along a gradient of increasing anthropogenic stress. Narrative rules for each of the BCG levels are presented to facilitate the evaluation of benthic communities in coral reefs and provide specific narrative features to detect changes in coral reef condition and biological integrity. The BCG model can be used in the absence of numeric, or quantitative metrics, to evaluate actions that may encroach on coral reef ecosystems, manage endangered species habitat, and develop and implement management plans for marine protected areas, watersheds, and coastal zones. The narrative BCG model is a defensible model and communication tool that translates scientific results so the nontechnical person can understand and support both regulatory and non-regulatory water quality and natural resource programs.

Puerto Rico, U.S. Virgin Islands

Climate change as a long-term stressor for the fisheries of the Laurentian Great Lakes of North America

The Laurentian Great Lakes of North America provide valuable ecosystem services, including fisheries, to the surrounding population. Given the prevalence of other anthropogenic stressors that have historically affected the fisheries of the Great Lakes (e.g., eutrophication, invasive species, overfishing), climate change is often viewed as a long-term stressor and, subsequently, may not always be prioritized by managers and researchers. However, climate change has the potential to negatively affect fish and fisheries in the Great Lakes through its influence on habitat. In this paper, we (1) summarize projected changes in climate and fish habitat in the Great Lakes; (2) summarize fish responses to climate change in the Great Lakes; (3) describe key interactions between climate change and other stressors relevant to Great Lakes fish, and (4) summarize how climate change can be incorporated into fisheries management. In general, fish habitat is projected to be characterized by warmer temperatures throughout the water column, less ice cover, longer periods of stratification, and more frequent and widespread periods of bottom hypoxia in productive areas of the Great Lakes. Based solely on thermal habitat, fish populations theoretically could experience prolonged optimal growth environment within a changing climate, however, models that assess physical habitat influences at specific life stages convey a more complex picture. Looking at specific interactions with other stressors, climate change may exacerbate the negative impacts of both eutrophication and invasive species for fish habitat in the Great Lakes. Although expanding monitoring and research to consider climate change interactions with currently studied stressors, may offer managers the best opportunity to keep the valuable Great Lakes fisheries sustainable, this expansion is globally applicable for large lake ecosystem dealing with multiple stressors in the face of continued human-driven changes.

Great Lakes

Quantifying resilience

The biosphere is under unprecedented pressure, reflected in rapid changes in our global ecological, social, technological and economic systems. In many cases, ecological and social systems can adapt to these changes over time, but when a critical threshold is surpassed, a system under stress can undergo catastrophic change and reorganize into a different state. The concept of resilience, introduced more than 40 years ago in the ecological sciences, captures the behaviour of systems that can occur in alternative states. The original definition of resilience forwarded by Holling ( 1973 ) is still the most useful. It defines resilience as the amount of disturbance that a system can withstand before it shifts into an alternative stable state. The idea of alternative stable states has clear and profound implications for ecological management. Coral reefs, for example, are high-diversity systems that provide key ecosystem services such as fisheries and coastal protection. Human impacts are causing significant, ongoing reef degradation, and many reefs have shifted from coral- to algal-dominated states in response to anthropogenic pressures such as elevated water temperatures and overfishing. Understanding and differentiating between the factors that help maintain reefs in coral-dominated states vs. those that facilitate a shift to an undesired algal-dominated state is a critical step towards sound management and conservation of these, and other, important social&ndash;ecological systems. Resilience has gained popularity among both academicians and laypeople, as a term meant to describe a systems&rsquo; ability to withstand disturbance. Resilience has become a buzzword in the last decade, as shown by its increasing appearance in calls for research proposals and scientific citation data bases. The term resilience has in many cases lost the clarity of the original definition and in fact is frequently used in a manner in direct opposition to the original definition. Many current uses of the concept are loose and incorrect. The term is becoming increasingly used in a normative sense (Brand & Jax 2007 ), as if resilience were a desirable quality of systems. However, even systems in highly undesirable states, such as macro-algae dominated reefs, or city cores in poverty traps, may be highly resilient, which is to say they withstand attempts to transform them into different (desirable) states. Operationalizing the concept of resilience for application and management has been difficult. Misuse of the term can have significant negative impacts, because resilience is being used to help guide responses to natural disasters and to assess the sustainability of ecosystems and urban systems and has been driving international research priorities. Resilience has been argued to be a basic emergent property of systems, a process or a rate. We focus on the original concept as described by Holling, which is that of an emergent system property; when a system is in a desirable state and managers wish to enhance resilience, or when the system is in an undesirable state and managers wish to erode resilience and foster a transformation to an alternative state. Fostering or eroding resilience is a process. When a system is perturbed but resilience is not exceeded, then the recovery can be measured as a rate. Several frameworks to operationalize resilience have been proposed. A decade ago, a special feature focused on quantifying resilience was published in the journal Ecosystems (Carpenter, Westley & Turner 2005 ). The approach there was towards identifying surrogates of resilience, but few of the papers proposed quantifiable metrics. Consequently, many ecological resilience frameworks remain vague and difficult to quantify, a problem that this special feature aims to address. However, considerable progress has been made during the last decade (e.g. Pope, Allen & Angeler 2014 ). Although some argue that resilience is best kept as an unquantifiable, vague concept (Quinlan et al . 2016 ), to be useful for managers, there must be concrete guidance regarding how and what to manage and how to measure success (Garmestani, Allen & Benson 2013 ; Spears et al . 2015 ). Ideas such as &lsquo;resilience thinking&rsquo; have utility in helping stakeholders conceptualize their systems, but provide little guidance on how to make resilience useful for ecosystem management, other than suggesting an ambiguous, Goldilocks approach of being just right (e.g. diverse, but not too diverse; connected, but not too connected). Here, we clarify some prominent resilience terms and concepts, introduce and synthesize the papers in this special feature on quantifying resilience and identify core unanswered questions related to resilience.

Journal of Applied Ecology

Predicting effects of large-scale reforestation on native and exotic birds

Aim Ecological restoration is critical for recovering biodiversity and ecosystem services, yet designing interventions to achieve particular outcomes remains fraught with challenges. In the extensive regions where non-native species are firmly established, it is unlikely that historical conditions can be fully reinstated. To what degree, and how rapidly, can human-dominated areas be shifted via restoration into regimes that benefit target species, communities or processes? Location We explore this question in a >20-year-old reforestation effort underway at Hakalau Forest National Wildlife Refuge in montane Hawaii. This large-scale planting of Acacia koa trees is designed to secure populations of globally threatened bird species by transitioning the site rapidly from pasture to native forest. Methods We surveyed all forest birds in multiple corridors of young planted trees, remnant corridors of mature trees along gulches and at sites within mature forest. Using a Bayesian hierarchical approach, we identified which factors (distance from forest, habitat type and surrounding tree cover) had the most important influence on native and exotic bird abundance in the reforestation area. Results We found that 90% of native and exotic bird species responded quickly, occupying corridors of native trees approximately a decade after planting. However, native and exotic forest birds responded to markedly different characteristics of the reforested area. Native bird abundance was strongly predicted by proximity to mature forest and remnant corridors; conversely, exotic bird abundance was best predicted by overall tree cover throughout the area reforested. Main conclusions Our results demonstrate that large-scale tree planting in corridors adjacent to mature forest can catalyse rapid recovery (both increased abundance and expanded distribution) of forest birds and that it is possible to design reforestation to benefit native species in novel ecosystems.

Hawaii

Shellfish aquaculture farms as foraging habitat for nearshore fishes and crabs

Objective Oyster reefs across North America have declined precipitously over the past 140 years. In Washington State, Olympia oyster Ostrea lurida reefs historically provided water filtration and nearshore structural habitat for fishes and invertebrates, but this species is now functionally extinct across its historical range. In place of these naturally occurring reefs, shellfish farms consisting mainly of nonnative Pacific oysters Magallana gigas now occupy patches of nearshore habitat across Washington. These farms modify intertidal substrate by adding structural habitat via suspended oyster grow bags, predator exclusion nets, loose oyster beds, and other shellfish grow-out gear. As interest and investment in shellfish aquaculture have expanded both locally and globally, so has interest in how these farms modify intertidal habitat and whether the complex structure created by the shellfish and shellfish growing gear provides ecosystem services that are comparable to those of unfarmed areas, such as mudflats and eelgrass meadows. Methods In this study, we sought to quantify how shellfish farms are used as foraging habitat for several common nearshore species of fish and crabs in Puget Sound, Washington. We used direct observations of species-specific behaviors from underwater video to model how habitat type affected observed foraging rates. Result We obtained a total of 393 crab observations, 431 demersal fish observations, and 1856 pelagic fish observations across all seven farm sites. Several common species of pelagic fish (e.g., surfperch [Embiotocidae]) used aquaculture-growing gear more frequently than unfarmed areas as foraging habitat, but Metacarcinus spp. crabs displayed higher foraging frequency in unfarmed mudflats. Species groups such as sculpins (Cottidae) and small flatfish (Pleuronectidae) clearly used specific aquaculture-growing gear and mudflats in roughly equal proportion. Conclusion Our results indicate that shellfish farms within a larger nearshore habitat mosaic of eelgrass meadows, mudflats, bivalve aquaculture gear, and edge habitat can provide foraging habitat for several species of nearshore fish.

Washington

Divergent climate impacts on C3 versus C4 grasses imply widespread 21st century shifts in grassland functional composition

Aim Grasslands cover a third of Earth's landmass and provide critical ecosystem services. Anticipating how perennial C 3 (cool-season) and C 4 (warm-season) grasses respond to climate change will be key to predicting future composition and functioning of grasslands. Here, we evaluate environmental drivers of C 3 and C 4 perennial distributions and assess how C 3 and C 4 grass distributions shift in response to future climate change. Location Western United States. Methods We developed integrated species distribution models to identify climate and soil drivers of relative abundance of C 3 and C 4 perennial grasses. We then created projections of species abundances under future climate and evaluated when and where projected shifts in relative abundance were robust across climate models. Results Historically, C 3 grasses occupied areas with lower temperature and more variable precipitation regimes, while C 4 grasses occupied areas of higher temperature, greater temperature variability and greater warm-season precipitation. C 4 species also occupied narrower soil texture niches. In response to future climate change, C 3 grass abundance declined across 74% of areas, while C 4 abundance increased across 66% of areas. C 3 grasses expanded in mid- to higher-latitude areas with increasing temperature and decreasing seasonality of precipitation. In contrast, C 4 grasses increased in higher-latitude regions, but declined in lower-latitude, dryer regions. Results were surprisingly robust across climate scenarios, suggesting high confidence in the direction of these future changes. Main Conclusions Findings imply C 3 and C 4 perennial grasses will have highly divergent responses to climate change that may result in grassland functional compositional changes. Increasing temperatures and precipitation variability may favour some C 4 grasses, but C 4 habitat expansion may be constrained by soil conditions in western USA. Results provide actionable insights for anticipating the impacts of climate change on grass-dominated and co-dominated ecosystems and improving large-scale conservation and restoration efforts.

Diversity and Distributions