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Social–ecological mismatches create conservation challenges in introduced species management

Introduced species can have important effects on the component species and processes of native ecosystems. However, effective introduced species management can be complicated by technical and social challenges. We identify “social–ecological mismatches” (that is, differences between the scales and functioning of interacting social and ecological systems) as one such challenge. We present three case studies in which mismatches between the organization and functioning of key social and ecological systems have contributed to controversies and debates surrounding introduced species management and policy. We identify three common issues: social systems and cultures may adapt to a new species’ arrival at a different rate than ecosystems; ecological impacts can arise at one spatial scale while social impacts occur at another; and the effects of introduced species can spread widely, whereas management actions are constrained by organizational and/or political boundaries. We propose strategies for collaborative knowledge building and adaptive management that may help address these challenges.

Frontiers in Ecology and the Environment

Mitigating amphibian disease: strategies to maintain wild populations and control chytridiomycosis

Background Rescuing amphibian diversity is an achievable conservation challenge. Disease mitigation is one essential component of population management. Here we assess existing disease mitigation strategies, some in early experimental stages, which focus on the globally emerging chytrid fungus Batrachochytrium dendrobatidis . We discuss the precedent for each strategy in systems ranging from agriculture to human medicine, and the outlook for each strategy in terms of research needs and long-term potential. Results We find that the effects of exposure to Batrachochytrium dendrobatidis occur on a spectrum from transient commensal to lethal pathogen. Management priorities are divided between (1) halting pathogen spread and developing survival assurance colonies, and (2) prophylactic or remedial disease treatment. Epidemiological models of chytridiomycosis suggest that mitigation strategies can control disease without eliminating the pathogen. Ecological ethics guide wildlife disease research, but several ethical questions remain for managing disease in the field. Conclusions Because sustainable conservation of amphibians in nature is dependent on long-term population persistence and co-evolution with potentially lethal pathogens, we suggest that disease mitigation not focus exclusively on the elimination or containment of the pathogen, or on the captive breeding of amphibian hosts. Rather, successful disease mitigation must be context specific with epidemiologically informed strategies to manage already infected populations by decreasing pathogenicity and host susceptibility. We propose population level treatments based on three steps: first, identify mechanisms of disease suppression; second, parameterize epizootiological models of disease and population dynamics for testing under semi-natural conditions; and third, begin a process of adaptive management in field trials with natural populations.

Frontiers in Zoology

Integrating climate change, biological invasions, and infectious wildlife diseases

Climate change is likely to affect infectious diseases that are facilitated by biological invasions, with repercussions for wildlife conservation and zoonotic risks. Current invasion management and policy are underprepared for the future risks associated with such invasion-related wildlife diseases. By considering evidence from bioclimatology, invasion biology, and disease research, we illustrate how climate change is anticipated to affect disease agents (parasites and pathogens), hosts, and vectors across the different stages of invasions. We highlight the opportunity to integrate these disciplines to identify the effects of climate change on invasion related wildlife diseases. In addition, shifting to a proactive stance in implementing management and policy, such as by incorporating climate-change effects either into preventative and mitigation measures for biosecurity or with rapid response protocols to limit disease spread and impacts, could help to combat future ecological, economic, and human health risks stemming from invasion-related wildlife diseases.

Frontiers in Ecology and the Environment

Editorial: Contributions of behavior and physiology to conservation biology

Conservation biology is a rapidly evolving discipline, with its synthetic, multidisciplinary framework expanding extensively in recent years. Seemingly disparate disciplines, such as behavior and physiology, are being integrated into this discipline's growing portfolio, resulting in diverse tools that can help develop conservation solutions. Behavior and physiology have traditionally been considered separate fields, yet their integration can provide a more comprehensive approach to developing solutions to conservation and management problems. However, demonstrations are needed of how behavior and physiology—either separately or combined—have contributed to conservation success. Examining species' vulnerabilities to extinction through the lenses of behavior and physiology can provide insight into the mechanisms that drive population declines and extirpations. Our goal is to increase awareness of the benefit of combining behavioral and physiological tools to improve conservation management decisions. Such studies can also help strengthen the basis for evidence-based conservation which, in some cases, has been previously lacking. The diverse studies in our Research Topic illustrate key examples of ways that behavior and physiology can be incorporated into conservation biology. Three main themes emerged from the invited papers with respect to their relevance to conservation: (1) Stress physiology, (2) indicators of health and disease dynamics, (3) and movement ecology. But these themes were also intertwined, thus showing the importance of integrating multiple fields of research to successfully address questions about conservation biology. Two mini reviews discuss the importance of examining how stress physiology may affect individual fitness and capacity to cope with change which, ultimately, affects the resiliency of populations. Walls and Gabor, in their mini review, promoted combining studies of behavior and physiology to aid in developing conservation strategies for amphibians, which could provide conservation managers with workable solutions to global environmental change. Walls and Gabor also pointed out that studies of behavior are useful to understand how native amphibian species respond to invasive predators. This is supported by Roznik et al. who combined physiology and behavior for a more holistic understanding of the impacts of competition between native and invasive frogs. These authors found that invasive Cuban tree frogs outperformed two native species in jump length and speed, offsetting the costs of dehydration. This work also ties in well to a review by Joly, who indicated the importance of exploring movement ecology (on multiple spatial scales) of invasive species to provide insights for invasive species management. Walls and Gabor also emphasized that more studies on phenological shifts of species in response to recent climate change are needed, as explored by Bókony et al. with birds. These authors found that bird species varied their migratory behaviors in response to contemporary climate change which, in turn, influenced subsequent trends in population size.

Frontiers in Ecology and Evolution

Optimal allocation of law enforcement patrol effort to mitigate poaching activities

Poaching is a global problem causing the decline of species worldwide. Optimizing the efficiency of ranger patrols to deter poaching activity at the lowest possible cost is crucial for protecting species with limited resources. We applied decision analysis and spatial optimization algorithms to allocate efforts of ranger patrols throughout a national park. Our objective was to mitigate poaching activity at or below management risk targets for the lowest monetary cost. We examined this trade‐off by constructing a Pareto efficiency frontier using integer linear programming. We used data from a ranger‐based monitoring program in Nyungwe National Park, Rwanda. Our measure of poaching risk is based on dynamic occupancy models that account for imperfect detection of poaching activities. We found that in order to achieve a 5% reduction in poaching risk, 622 ranger patrol events (each corresponding to patrolling 1‐km 2 sites) were needed within a year at a cost of US$49,760. In order to attain a 60% reduction in poaching risk, 15,560 patrol events were needed at a cost of US \$ 1,244,800. We evaluated the trade‐off between patrol cost and poaching risk based on our model by constructing a Pareto efficiency frontier and park managers found the solution for a 50% risk reduction to be a practical trade‐off based on funding constraints (comparable to recent years) and the diminishing returns between risk mitigation and cost. This expected reduction in risk required 8,558 patrol events per year at a cost of US \$ 684,640. Our results suggest that optimal solutions could increase efficiency compared to the actual effort allocations from 2006 to 2016 in Nyungwe National Park (e.g., risk reductions of ~30% under recent budgets compared to ~50% reduction in risk under the optimal strategy). The modeling framework in this study took into account imperfect detection of poaching risk as well as the directional and conditional nature of ranger patrol events given the spatial adjacency relationships of neighboring sites and access points. Our analyses can help to improve the efficiency of ranger patrols, and the modeling framework can be broadly applied to other spatial conservation planning problems with conditional, multilevel, site selection.

Nyungwe National Park

A call for conservation scientists to evaluate opportunities and risks from operation of vertical axis wind turbines

A new conservation paradigm ( Kareiva and Marvier, 2012 ) emphasizes the need for scientists to embrace a holistic approach taking into account the social and natural dimensions of conservation in human-dominated landscapes. While there is heavy debate over the new approach ( Tallis and Lubchenco, 2014 ), most conservation scientists seem to agree on to the need to cooperate with corporations when such interaction can benefit people and the environment ( Miller et al., 2014 ; Tallis and Lubchenco, 2014 ). Cooperation can be most productive when established in the early phases of development, but this requires a high capacity for forward looking pre-emptive action (i.e., anticipating potential forthcoming issues before they arise; Sutherland and Woodroof, 2009 ). This framework is particularly salient for rapidly developing and expanding technologies such as those for harvesting renewable energy sources. Here the stakes are very high, as they concern mitigating negative consequences to global climate while generating energy without impacting wildlife. In this vein, past experience is instructional. The environmental impacts of biofuels and wind, among others, have been identified and evaluated rather late ( Sutherland and Woodroof, 2009 ), so that implementation of best management practices on existing facilities is now often prohibitively expensive.

Frontiers in Ecology and Evolution

Diverse portfolios: Investing in tributaries for restoration of large river fishes in the Anthropocene

Rehabilitation of large Anthropocene rivers requires engagement of diverse stakeholders across a broad range of sociopolitical boundaries. Competing objectives often constrain options for ecological restoration of large rivers whereas fewer competing objectives may exist in a subset of tributaries. Further, tributaries contribute toward building a “portfolio” of river ecosystem assets through physical and biological processes that may present opportunities to enhance the resilience of large river fishes. Our goal is to review roles of tributaries in enhancing mainstem large river fish populations. We present case histories from two greatly altered and distinct large-river tributary systems that highlight how tributaries contribute four portfolio assets to support large-river fish populations: 1) habitat diversity, 2) connectivity, 3) ecological asynchrony, and 4) density-dependent processes. Finally, we identify future research directions to advance our understanding of tributary roles and inform conservation actions. In the Missouri River United States, we focus on conservation efforts for the state endangered lake sturgeon, which inhabits large rivers and tributaries in the Midwest and Eastern United States. In the Colorado River, Grand Canyon United States, we focus on conservation efforts for recovery of the federally threatened humpback chub. In the Missouri River, habitat diversity focused on physical habitats such as substrate for reproduction, and deep-water habitats for refuge, whereas augmenting habitat diversity for Colorado River fishes focused on managing populations in tributaries with minimally impaired thermal and flow regimes. Connectivity enhancements in the Missouri River focused on increasing habitat accessibility that may require removal of physical structures like low-head dams; whereas in the Colorado River, the lack of connectivity may benefit native fishes as the disconnection provides refuge from non-native fish predation. Hydrologic variability among tributaries was present in both systems, likely underscoring ecological asynchrony. These case studies also described density dependent processes that could influence success of restoration actions. Although actions to restore populations varied by river system, these examples show that these four portfolio assets can help guide restoration activities across a diverse range of mainstem rivers and their tributaries. Using these assets as a guide, we suggest these can be transferable to other large river-tributary systems.

Frontiers in Environmental Science

Can polar bears use terrestrial foods to offset lost ice-based hunting opportunities?

Increased land use by polar bears ( Ursus maritimus ) due to climate-change-induced reduction of their sea-ice habitat illustrates the impact of climate change on species distributions and the difficulty of conserving a large, highly specialized carnivore in the face of this global threat. Some authors have suggested that terrestrial food consumption by polar bears will help them withstand sea-ice loss as they are forced to spend increasing amounts of time on land. Here, we evaluate the nutritional needs of polar bears as well as the physiological and environmental constraints that shape their use of terrestrial ecosystems. Only small numbers of polar bears have been documented consuming terrestrial foods even in modest quantities. Over much of the polar bear's range, limited terrestrial food availability supports only low densities of much smaller, resident brown bears ( Ursus arctos ), which use low-quality resources more efficiently and may compete with polar bears in these areas. Where consumption of terrestrial foods has been documented, polar bear body condition and survival rates have declined even as land use has increased. Thus far, observed consumption of terrestrial food by polar bears has been insufficient to offset lost ice-based hunting opportunities but can have ecological consequences for other species. Warming-induced loss of sea ice remains the primary threat faced by polar bears.

Frontiers in Ecology and the Environment

Temporal coherence patterns of prairie pothole wetlands indicate the importance of landscape linkages and wetland heterogeneity in maintaining biodiversity

Wetland ecosystems are diverse, productive habitats that are essential reservoirs of biodiversity. Not only are they home to numerous wetland-specialist species, but they also provide food, water, and shelter that support terrestrial wildlife populations. However, like observed patterns of biodiversity loss, wetland habitats have experienced widespread loss and degradation. In order to conserve and restore wetlands, and thereby the biodiversity they support, it is important to understand how biodiversity in wetland habitats is maintained. Habitat heterogeneity and connectivity are thought to be predominate drivers of wetland biodiversity. We quantified temporal coherence (i.e., spatial synchrony) of wetland invertebrate communities using intra-class correlations among 16 wetlands sampled continuously over 24 years to better understand the relative influences wetland heterogeneity (i.e., internal processes specific to individual wetlands and spatial connectivity and external processes occurring on the landscape) on wetland biodiversity. We found that while wetlands with different ponded-water regimes (temporarily ponded or permanently ponded) often hosted different invertebrate communities, temporal shifts in invertebrate composition were synchronous. We also found the relative importance of internal versus external forces in determining community assembly vary depending on a wetland’s hydrologic function and climate influences. Our results confirm that heterogeneity and spatial connectivity of wetland landscapes are important drivers of wetland biodiversity.

North Dakota

A novel framework to predict relative habitat selection in aquatic systems: Applying machine learning and resource selection functions to acoustic telemetry data from multiple shark species

Resource selection functions (RSFs) have been widely applied to animal tracking data to examine relative habitat selection and to help guide management and conservation strategies. While readily used in terrestrial ecology, RSFs have yet to be extensively used within marine systems. As acoustic telemetry continues to be a pervasive approach within marine environments, incorporation of RSFs can provide new insights to help prioritize habitat protection and restoration to meet conservation goals. To overcome statistical hurdles and achieve high prediction accuracy, machine learning algorithms could be paired with RSFs to predict relative habitat selection for a species within and even outside the monitoring range of acoustic receiver arrays, making this a valuable tool for marine ecologists and resource managers. Here, we apply RSFs using machine learning to an acoustic telemetry dataset of four shark species to explore and predict species-specific habitat selection within a marine protected area. In addition, we also apply this RSF-machine learning approach to investigate predator-prey relationships by comparing and averaging tiger shark relative selection values with the relative selection values derived for eight potential prey-species. We provide methodological considerations along with a framework and flexible approach to apply RSFs with machine learning algorithms to acoustic telemetry data and suggest marine ecologists and resource managers consider adopting such tools to help guide both conservation and management strategies.

Buck Island Reef National Monument, St Croix

Combining physical and species‐based approaches improves refugia identification

Climate‐change refugia – locations likely to facilitate species persistence under climate change – are increasingly important components of conservation planning. Recent approaches for identifying refugia at broad scales include identifying regions that are projected to experience less severe changes (climatic exposure), that contain a diversity of physical and topographic features (environmental diversity), and that either retain or remain close to suitable climatic conditions (climate tracking, including both “species‐neutral” and species‐based approaches). We compared the degree of agreement between these approaches – with respect to their spatial coverage and other characteristics – across much of North America. This analysis found that approaches based on environmental diversity and species‐neutral climatic gradients both favored topographically complex regions, whereas climatic exposure and species‐based approaches identified regions with a range of topographic characteristics. Species‐based approaches targeting specific habitat groups identified unique regions missed by other approaches, emphasizing the importance of asking the question “refugia for what?” when prioritizing refugia. Our results highlight the necessity of including climatic exposure and species‐based information in addition to topographic diversity and climatic gradients in refugia analyses.

Frontiers in Ecology and the Environment

Establishing the foundation for the global observing system for marine life

Maintaining healthy, productive ecosystems in the face of pervasive and accelerating human impacts including climate change requires globally coordinated and sustained observations of marine biodiversity. Global coordination is predicated on an understanding of the scope and capacity of existing monitoring programs, and the extent to which they use standardized, interoperable practices for data management. Global coordination also requires identification of gaps in spatial and ecosystem coverage, and how these gaps correspond to management priorities and information needs. We undertook such an assessment by conducting an audit and gap analysis from global databases and structured surveys of experts. Of 371 survey respondents, 203 active, long-term (>5 years) observing programs systematically sampled marine life. These programs spanned about 7% of the ocean surface area, mostly concentrated in coastal regions of the United States, Canada, Europe, and Australia. Seagrasses, mangroves, hard corals, and macroalgae were sampled in 6% of the entire global coastal zone. Two-thirds of all observing programs offered accessible data, but methods and conditions for access were highly variable. Our assessment indicates that the global observing system is largely uncoordinated which results in a failure to deliver critical information required for informed decision-making such as, status and trends, for the conservation and sustainability of marine ecosystems and provision of ecosystem services. Based on our study, we suggest four key steps that can increase the sustainability, connectivity and spatial coverage of biological Essential Ocean Variables in the global ocean: (1) sustaining existing observing programs and encouraging coordination among these; (2) continuing to strive for data strategies that follow FAIR principles (findable, accessible, interoperable, and reusable); (3) utilizing existing ocean observing platforms and enhancing support to expand observing along coasts of developing countries, in deep ocean basins, and near the poles; and (4) targeting capacity building efforts. Following these suggestions could help create a coordinated marine biodiversity observing system enabling ecological forecasting and better planning for a sustainable use of ocean resources.

Frontiers in Marine Science

Modeling functional flows in California rivers

Environmental flows are critical to the recovery and conservation of freshwater ecosystems worldwide. However, estimating desired ranges of environmental flows across large, diverse landscapes is challenging. To advance protections of environmental flows for streams in California, USA, we developed a statewide modeling approach focused on functional components of the natural flow regime. Functional flow components in California streams—fall pulse flows, wet season peak flows and base flows, the spring flow recession, and dry season baseflows—support essential physical and ecological processes in riverine ecosystems. These functional flow components can be represented by functional flow metrics (FFMs) and quantified by their magnitude, timing, frequency, duration, and rate-of-change from daily streamflow records. After quantifying FFMs at reference-quality streamflow gages in California, we used machine-learning methods to estimate their natural range of values for all stream reaches in the state based on physical watershed characteristics and climatic factors. We found that the models performed well in predicting FFMs in streams across a diversity of landscape and climate contexts, according to several model performance criteria. Using the predicted FFM values, we established initial estimates of ecological flows that are expected to support critical functions and are broadly protective of ecosystem health. Modeling functional flows statewide offers a pathway for increasing the pace and scale of environmental flow protections in California and beyond.

California

Adult monarch (Danaus plexippus) abundance is higher in burned sites than in grazed sites

Much of the remaining suitable habitat for monarchs (Danaus plexippus) in Minnesota is found in tallgrass prairies. We studied the association of adult monarch abundance with use of fire or grazing to manage prairies. Sites (n=20) ranged in size from 1 to 145 hectares and included land owned and managed by the Minnesota DNR, U.S. Fish and Wildlife Service, The Nature Conservancy, and private landowners. We measured Asclepias spp. (milkweeds, monarch host plants) and forb frequency in 0.5 x 2-m plots located along randomly-placed transects that were stratified to sample wet, mesic, and dry prairie types at each site. Adult butterfly surveys took place three times at each site during the summers of 2016 and 2017, using a standardized Pollard Walk (400 meters). Data were analyzed using mixed effects models. Monarchs were more abundant at sites managed with prescribed fire than with grazing. We found no difference in milkweed and forb frequency between burned and grazed prairies. There was no relationship between monarch abundance and the other predictor variables tested: milkweed frequency, site area, forb frequency, and percent prairie in a 1.5 km buffer area surrounding each site. Monarch abundance was lowest at grazed sites with high stocking rates. Our findings suggest that the use of burning or grazing for prairie management is not associated with milkweed or forb frequency, at least for sites that have not been burned in several years. They also suggest that heavy grazing may have negative impacts on monarchs.

Minnesota

Wet meadow regeneration through restoration of biophysical feedbacks

Wet meadows are globally significant ecosystems that provide critical hydrological, ecological, and biogeochemical functions, yet their extent has declined dramatically due to land use changes and hydrologic alteration. These sedge-dominated wetlands exist at the drier end of the wetland gradient, maintained by shallow groundwater and periodic inundation. This paper is a global synthesis of the ecological, geomorphic, and hydrological dynamics of wet meadows, with an emphasis on alluvial systems, to inform effective restoration strategies. We compare wet meadows to other wetlands, classify them into palustrine, lacustrine, and alluvial types, then focus on alluvial wet meadows and discuss how their formation and persistence depend on ground and surface water interactions, sediment deposition and flow obstructions, all mediated by biological processes. In particular, we highlight the role of hydric graminoids in resisting erosion and maintaining soil cohesion, how beaver promote meadow persistence, and the significance of wet meadows as carbon sinks. We also present stratigraphic evidence demonstrating that incision, often triggered by anthropogenic activity or changing climate, is the primary mechanism of alluvial wet meadow degradation, resulting in water table decline and shifts in vegetation composition. Restoration requires reversing these incisional processes through techniques that elevate water tables, disperse flow and retain sediment—methods traditionally associated with either soil conservation or stream restoration. These include nature-based solutions that create obstructions such as beaver dams and their analogues, rock and wood-based obstructions and incision trench or gully filling and grading. Given their multifunctional value—including but not limited to flood attenuation, biodiversity support, and carbon sequestration—wet meadows warrant a focused restoration framework. This review advocates for a valley-floor scale restoration paradigm that integrates hydrological reconnection, sediment retention, and biological reinforcement to ensure long-term resilience of these systems in the face of changing climate and land use pressures.

Frontiers in Environmental Science

Variability in coastal habitat available for Longfin Smelt Spirinchus thaleichthys in the northeastern Pacific Ocean

An understanding of oceanographic conditions and processes important to marine animal ecology is fundamental to the development of effective management and conservation actions. Longfin Smelt ( Spirinchus thaleichthys ) is a pelagic forage fish found in coastal and estuarine waters along the Pacific coast of North America from Alaska to central California. Substantial population declines in California’s San Francisco Estuary, where Longfin Smelt are protected under California’s Endangered Species Act, have prompted extensive study of estuarine factors associated with the decline. However, coastal factors that affect up to two-thirds of the Longfin Smelt life cycle are poorly understood and may be important drivers of population dynamics. We compiled coastal observations from numerous sources to estimate the range-wide coastal marine distribution of Longfin Smelt and assess habitat factors affecting distribution in the northeast Pacific Ocean. Based on maximum entropy species distribution models, Longfin Smelt distribution was correlated with depth, distance from the nearest estuary, sea surface temperature, and sea surface chlorophyll. Longfin Smelt were found in shallow, higher productivity coastal waters closer to estuaries, with depth and temperature the most consistent factors influencing distribution. Habitat suitability was highly variable at the southern extent of the range, particularly off the California coast, and was largely driven by habitat contractions associated with warm-water conditions. Study results provide insights into the habitat and range-wide distribution of an at-risk estuarine-reliant forage fish and are the first step toward identifying processes that affect the marine portion of the Longfin Smelt life cycle.

Frontiers in Marine Science

Mass mortality of avian migrants in New Mexico, USA, that coincided with an extreme weather event

Many birds are migratory, and this life history strategy allows for maximized access to seasonally abundant resources and favorable climates. However, migration exposes birds to threats and stressors, resulting in high mortality during migration. Anthropogenic landscape alterations and climate change have intensified threats, and mass mortality events linked to extreme weather are more common in recent decades. Documenting mass mortality is critical for predicting future occurrences and implementing effective conservation. Here, we describe a mortality event that occurred throughout New Mexico, USA in fall 2020. Carcasses began appearing in the region in mid-August, during a period of extreme heat and drought. Following an extreme cold weather event on September 8–9 th , the number of carcasses increased dramatically and expanded throughout the state. In total, we collected 628 carcasses comprising 58 species within Doña Ana and Otero Counties in New Mexico. Necropsy determined emaciation was the cause of death for 74.6% of carcasses. Live birds captured during the period of peak mortality (n = 223) were in similarly poor condition. This event provides a striking example of how multiple types of extreme stressors, in this case widespread drought and unseasonal cold, coincided with a mortality event, indicating a possible synergistic relationship between these factors and the mass mortality. Mortality events are likely to increase in frequency with intensifying climate change. Establishment of networks of biologists and researchers could improve our ability to identify and communicate developing mortality events, organize data collection, and improve understanding of the causes and consequences of mortality.

New Mexico

Shorebird reproductive response to exceptionally early and late springs varies across sites in Arctic Alaska

While increases in overall temperatures are widely reported in the Arctic, large inter-annual variation in spring weather, with extreme early and late conditions, is also occurring. Using data collected from three sites in Arctic Alaska, we explored how shorebird breeding density, nest initiation, nest synchrony, nest survival, and phenological mismatch varied between two exceptionally early (2015 and 2016) and late (2017 and 2018) springs. We assessed these differences in the context of long-term data from each site and whether species exhibited conservative or opportunistic reproductive strategies. Conservative shorebirds typically display nest-site fidelity and territoriality, consistent population densities, relatively even individual spacing, and monogamous mating systems with bi-parental incubation. In contrast, opportunistic shorebirds display the opposite traits, and a polygamous mating system with uniparental incubation. In this study, we evaluated 2,239 nests from 13 shorebird species, 2015–2018, and found that shorebirds of both strategies bred earlier and in higher numbers in early, warm springs relative to historic levels (based on 3,789 nests, 2005–2014); opposite trends were observed in late springs. In early springs, nests were initiated less synchronously than in late springs. Nest survival was unrelated to spring type, but was greater in earlier laid nests overall. Invertebrate food resources emerged earlier in early springs, resulting in a greater temporal asynchrony between invertebrate emergence and chick hatching in early than late springs. However, invertebrate abundance was quite variable among sites and years regardless of spring type. Overall, our results were generally consistent with predicted relationships between spring conditions and reproductive parameters. However, we detected differences among sites that could not be explained by other ecological factors (e.g., predators or alternative prey). Differences in shorebird community composition and other subtler methodological/ecological differences among sites highlight the difficulty of understanding the complex nature of these ecological systems and the importance of evaluating questions at multiple sites across multiple years. Our study demonstrates that shorebirds exhibit a high degree of behavioral flexibility in response to variable Arctic conditions, but whether this flexibility is enough to allow them to optimally track changing environmental conditions or if evolutionary adjustments will be necessary is unknown.

Alaska