Limited shifts in the distribution of migratory bird breeding habitat density in response to future changes in climate.

Grasslands, and the depressional wetlands that exist throughout them, are endangered ecosystems that face both climate and land-use change pressures. Tens of millions of dollars are invested annually to manage the existing fragments of these ecosystems to serve as critical breeding habitat for migratory birds. The North American Prairie Pothole Region (PPR) contains millions of depressional wetlands that produce between 50% and 80% of the continent's waterfowl population. Previous modeling efforts suggested that climate change would result in a shift of suitable waterfowl breeding habitat from the central to the southeast portion of the PPR, an area where over half of the depressional wetlands have been drained. The implications of these projections suggest a massive investment in wetland restoration in the southeastern PPR would be needed to sustain waterfowl populations at harvestable levels. We revisited these modeled results indicating how future climate may impact the distribution of waterfowl-breeding habitat using up-to-date climate model projections and a newly developed model for simulating prairie-pothole wetland hydrology. We also presented changes to the number of "May ponds," a metric used by the U.S. Fish and Wildlife Service to estimate waterfowl breeding populations and establish harvest regulations. Based on the output of 32 climate models and two emission scenarios, we found no evidence that the distribution of May ponds would shift in the future. However, our results projected a 12% decrease to 1% increase in May pond numbers when comparing the most recent climate period (1989-2018) to the end of the 21st century (2070-2099). When combined, our results suggest areas in the PPR that currently support the highest densities of intact wetland basins, and thus support the largest numbers of breeding-duck pairs, will likely also be the places most critical to maintaining continental waterfowl populations in an uncertain future.

Climate- versus geographic-dependent patterns in the spatial distribution of macroinvertebrate assemblages in New World depressional wetlands.

Analyses of biota at lower latitudes may presage impacts of climate change on biota at higher latitudes. Macroinvertebrate assemblages in depressional wetlands may be especially sensitive to climate change because weather-related precipitation and evapotranspiration are dominant ecological controls on habitats, and organisms of depressional wetlands are temperature-sensitive ectotherms. We aimed to better understand how wetland macroinvertebrate assemblages were structured according to geography and climate. To do so, we contrasted aquatic-macroinvertebrate assemblage structure (family level) between subtropical and temperate depressional wetlands of North and South America using presence-absence data from 264 of these habitats across the continents and more-detailed relative-abundance data from 56 depressional wetlands from four case-study locations (North Dakota and Georgia in North America; southern Brazil and Argentinian Patagonia in South America). Both data sets roughly partitioned wetland numbers equally between the two climatic zones and between the continents. We used ordination methods (PCA and NMDS) and tests of multivariate dispersion (PERMDISP) to assess the distribution and the homogeneity in variation in the composition of macroinvertebrate assemblages across climates and continents, respectively. We found that macroinvertebrate assemblage structures in the subtropical depressional wetlands of North and South America were similar to each other (at the family level), while assemblages in the North and South American temperate wetlands were unique from the subtropics, and from each other. Tests of homogeneity of multivariate dispersion indicated that family-level assemblage structures were more homogeneous in wetlands from the subtropical than the temperate zones. Our study suggests that ongoing climate change may result in the homogenization of macroinvertebrate assemblage structures in temperate zones of North and South America, with those assemblages becoming enveloped by assemblages from the subtropics. Biotic homogenization, more typically associated with other kinds of anthropogenic factors, may also be affected by climate change.

Do geographically isolated wetlands influence landscape functions?

Geographically isolated wetlands (GIWs), those surrounded by uplands, exchange materials, energy, and organisms with other elements in hydrological and habitat networks, contributing to landscape functions, such as flow generation, nutrient and sediment retention, and biodiversity support. GIWs constitute most of the wetlands in many North American landscapes, provide a disproportionately large fraction of wetland edges where many functions are enhanced, and form complexes with other water bodies to create spatial and temporal heterogeneity in the timing, flow paths, and magnitude of network connectivity. These attributes signal a critical role for GIWs in sustaining a portfolio of landscape functions, but legal protections remain weak despite preferential loss from many landscapes. GIWs lack persistent surface water connections, but this condition does not imply the absence of hydrological, biogeochemical, and biological exchanges with nearby and downstream waters. Although hydrological and biogeochemical connectivity is often episodic or slow (e.g., via groundwater), hydrologic continuity and limited evaporative solute enrichment suggest both flow generation and solute and sediment retention. Similarly, whereas biological connectivity usually requires overland dispersal, numerous organisms, including many rare or threatened species, use both GIWs and downstream waters at different times or life stages, suggesting that GIWs are critical elements of landscape habitat mosaics. Indeed, weaker hydrologic connectivity with downstream waters and constrained biological connectivity with other landscape elements are precisely what enhances some GIW functions and enables others. Based on analysis of wetland geography and synthesis of wetland functions, we argue that sustaining landscape functions requires conserving the entire continuum of wetland connectivity, including GIWs.

Differing Modes of Biotic Connectivity within Freshwater Ecosystem Mosaics.

We describe a collection of aquatic and wetland habitats in an inland landscape, and their occurrence within a terrestrial matrix, as a "freshwater ecosystem mosaic" (FEM). Aquatic and wetland habitats in any FEM can vary widely, from permanently ponded lakes, to ephemerally ponded wetlands, to groundwater-fed springs, to flowing rivers and streams. The terrestrial matrix can also vary, including in its influence on flows of energy, materials, and organisms among ecosystems. Biota occurring in a specific region are adapted to the unique opportunities and challenges presented by spatial and temporal patterns of habitat types inherent to each FEM. To persist in any given landscape, most species move to recolonize habitats and maintain mixtures of genetic materials. Species also connect habitats through time if they possess needed morphological, physiological, or behavioral traits to persist in a habitat through periods of unfavorable environmental conditions. By examining key spatial and temporal patterns underlying FEMs, and species-specific adaptations to these patterns, a better understanding of the structural and functional connectivity of a landscape can be obtained. Fully including aquatic, wetland, and terrestrial habitats in FEMs facilitates adoption of the next generation of individual-based models that integrate the principles of population, community, and ecosystem ecology.

The Significant Surface-Water Connectivity of "Geographically Isolated Wetlands".

We evaluated the current literature, coupled with our collective research expertise, on surface-water connectivity of wetlands considered to be "geographically isolated" (sensu Tiner Wetlands 23:494-516, 2003a) to critically assess the scientific foundation of grouping wetlands based on the singular condition of being surrounded by uplands. The most recent research on wetlands considered to be "geographically isolated" shows the difficulties in grouping an ecological resource that does not reliably indicate lack of surface water connectivity in order to meet legal, regulatory, or scientific needs. Additionally, the practice of identifying "geographically isolated wetlands" based on distance from a stream can result in gross overestimates of the number of wetlands lacking ecologically important surface-water connections. Our findings do not support use of the overly simplistic label of "geographically isolated wetlands". Wetlands surrounded by uplands vary in function and surface-water connections based on wetland landscape setting, context, climate, and geographic region and should be evaluated as such. We found that the "geographically isolated" grouping does not reflect our understanding of the hydrologic variability of these wetlands and hence does not benefit conservation of the Nation's diverse wetland resources. Therefore, we strongly discourage use of categorizations that provide overly simplistic views of surface-water connectivity of wetlands fully embedded in upland landscapes.

Using a vegetation index to assess wetland condition in the Prairie Pothole Region of North America.

Wetlands deliver a suite of ecosystem services to society. Anthropogenic activities, such as wetland drainage, have resulted in considerable wetland loss and degradation, diminishing the intrinsic value of wetland ecosystems worldwide. Protecting remaining wetlands and restoring degraded wetlands are common management practices to preserve and reclaim wetland benefits to society. Accordingly, methods for monitoring and assessing wetlands are required to evaluate their ecologic condition and outcomes of restoration activities. We used an established methodology for conducting vegetation-based assessments and describe a case study consisting of a wetland condition assessment in the Prairie Pothole Region of the North American Great Plains. We provide an overview of an existing method for selecting wetlands to sample across broad geographic distributions using a spatially balanced statistical design. We also describe site assessment protocols, including vegetation survey methods, and how field data were applied to a vegetation index that categorized wetlands according to ecologic condition. Results of the case study indicated that vegetation communities in nearly 50% of the surveyed wetlands were in very poor or poor condition, while only about 25% were considered good or very good. Approximately 70% of wetlands in native grasslands were categorized as good or very good compared to only 12% of those in reseeded grasslands (formerly cropland). In terms of informing restoration and management activities, results indicated that improved restoration practices could include a greater focus on establishing natural vegetation communities, and both restored and native prairie wetlands would benefit from enhanced management of invasive species.

Vulnerable Waters are Essential to Watershed Resilience.

Watershed resilience is the ability of a watershed to maintain its characteristic system state while concurrently resisting, adapting to, and reorganizing after hydrological (for example, drought, flooding) or biogeochemical (for example, excessive nutrient) disturbances. Vulnerable waters include non-floodplain wetlands and headwater streams, abundant watershed components representing the most distal extent of the freshwater aquatic network. Vulnerable waters are hydrologically dynamic and biogeochemically reactive aquatic systems, storing, processing, and releasing water and entrained (that is, dissolved and particulate) materials along expanding and contracting aquatic networks. The hydrological and biogeochemical functions emerging from these processes affect the magnitude, frequency, timing, duration, storage, and rate of change of material and energy fluxes among watershed components and to downstream waters, thereby maintaining watershed states and imparting watershed resilience. We present here a conceptual framework for understanding how vulnerable waters confer watershed resilience. We demonstrate how individual and cumulative vulnerable-water modifications (for example, reduced extent, altered connectivity) affect watershed-scale hydrological and biogeochemical disturbance response and recovery, which decreases watershed resilience and can trigger transitions across thresholds to alternative watershed states (for example, states conducive to increased flood frequency or nutrient concentrations). We subsequently describe how resilient watersheds require spatial heterogeneity and temporal variability in hydrological and biogeochemical interactions between terrestrial systems and down-gradient waters, which necessitates attention to the conservation and restoration of vulnerable waters and their downstream connectivity gradients. To conclude, we provide actionable principles for resilient watersheds and articulate research needs to further watershed resilience science and vulnerable-water management.

Cross-Ecosystem Fluxes of Pesticides from Prairie Wetlands Mediated by Aquatic Insect Emergence: Implications for Terrestrial Insectivores.

Contaminants alter the quantity and quality of insect prey available to terrestrial insectivores. In agricultural regions, the quantity of aquatic insects emerging from freshwaters can be impacted by insecticides originating from surrounding croplands. We hypothesized that, in such regions, adult aquatic insects could also act as vectors of pesticide transfer to terrestrial food webs. To estimate insect-mediated pesticide flux from wetlands embedded in an important agricultural landscape, semipermanetly and temporarily ponded wetlands were surveyed in cropland and grassland landscapes across a natural salinity gradient in the Prairie Pothole Region of North Dakota (USA) during the bird breeding season in 2015 and 2016 (n = 14 and 15 wetlands, respectively). Current-use pesticides, including the herbicide atrazine and the insecticides bifenthrin and imidacloprid, were detected in newly emerged insects. Pesticide detections were similar in insects emerging from agricultural and grassland wetlands. Biomass of emerging aquatic insects decreased 43%, and insect-mediated pesticide flux increased 50% along the observed gradient in concentrations of insecticides in emerging aquatic insects (from 3 to 577 ng total insecticide g-1 insect). Overall, adult aquatic insects were estimated to transfer between 2 and 180 µg total pesticide wetland-1 d-1 to the terrestrial ecosystem. In one of the 2 study years, biomass of emerging adult aquatic insects was also 73% lower from agricultural than grassland wetlands and was dependent on salinity. Our results suggest that accumulated insecticides reduce the availability of adult aquatic insect prey for insectivores and potentially increase insectivore exposure to insect-borne pesticides. Adult aquatic insects retain pesticides across metamorphosis and may expose insectivores living near both agricultural and grassland wetlands to dietary sources of toxic chemicals. Environ Toxicol Chem 2021;40:2282-2296. Published 2021. This article is a U.S. Government work and is in the public domain in the USA.

Alternative stable states in inherently unstable systems.

Alternative stable states are nontransitory states within which communities can exist. However, even highly dynamic communities can be viewed within the framework of stable-state theory if an appropriate "ecologically relevant" time scale is identified. The ecologically relevant time scale for dynamic systems needs to conform to the amount of time needed for a system's community to complete an entire cycle through its normal range of variation. For some systems, the ecologically relevant period can be relatively short (eg, tidal systems), for others it can be decadal (eg, prairie wetlands). We explore the concept of alternative stable states in unstable systems using the highly dynamic wetland ecosystems of North America's Prairie Pothole Region. The communities in these wetland ecosystems transition through multiple states in response to decadal-long climate oscillations that cyclically influence ponded-water depth, permanence, and chemistry. The perspective gained by considering dynamic systems in the context of stable-state theory allows for an increased understanding of how these systems respond to changing drivers that can push them past tipping points into alternative states. Incorporation of concepts inherent to stable-state theory has been suggested as a key scientific element upon which to base sustainable environmental management.

Modeling effects of crop production, energy development and conservation-grassland loss on avian habitat.

Birds are essential components of most ecosystems and provide many services valued by society. However, many populations have undergone striking declines as their habitats have been lost or degraded by human activities. Terrestrial grasslands are vital habitat for birds in the North American Prairie Pothole Region (PPR), but grassland conversion and fragmentation from agriculture and energy-production activities have destroyed or degraded millions of hectares. Conservation grasslands can provide alternate habitat. In the United States, the Conservation Reserve Program (CRP) is the largest program maintaining conservation grasslands on agricultural lands, but conservation grasslands in the PPR have declined by over 1 million ha since the program's zenith in 2007. We used an ecosystem-services model (InVEST) parameterized for the PPR to quantify grassland-bird habitat remaining in 2014 and to assess the degradation status of the remaining grassland-bird habitat as influenced by crop and energy (i.e., oil, natural gas, and wind) production. We compared our resultant habitat-quality ratings to grassland-bird abundance data from the North American Breeding Bird Survey to confirm that ratings were related to grassland-bird abundance. Of the grassland-bird habitat remaining in 2014, about 19% was degraded by crop production that occurred within 0.1 km of grassland habitats, whereas energy production degraded an additional 16%. We further quantified the changes in availability of grassland-bird habitat under various land-cover scenarios representing incremental losses (10%, 25%, 50%, 75%, and 100%) of CRP grasslands from 2014 levels. Our model identified 1 million ha (9%) of remaining grassland-bird habitat in the PPR that would be lost or degraded if all CRP conservation grasslands were returned to crop production. Grassland regions world-wide face similar challenges in maintaining avian habitat in the face of increasing commodity and energy production to sate the food and energy needs of a growing world population. Identifying ways to model the impacts of the tradeoff between food and energy production and wildlife production is an important step in creating solutions.

Landscape genetics reveal broad and fine-scale population structure due to landscape features and climate history in the northern leopard frog (Rana pipiens) in North Dakota.

Prehistoric climate and landscape features play large roles structuring wildlife populations. The amphibians of the northern Great Plains of North America present an opportunity to investigate how these factors affect colonization, migration, and current population genetic structure. This study used 11 microsatellite loci to genotype 1,230 northern leopard frogs (Rana pipiens) from 41 wetlands (30 samples/wetland) across North Dakota. Genetic structure of the sampled frogs was evaluated using Bayesian and multivariate clustering methods. All analyses produced concordant results, identifying a major east-west split between two R. pipiens population clusters separated by the Missouri River. Substructuring within the two major identified population clusters was also found. Spatial principal component analysis (sPCA) and variance partitioning analysis identified distance, river basins, and the Missouri River as the most important landscape factors differentiating R. pipiens populations across the state. Bayesian reconstruction of coalescence times suggested the major east-west split occurred ~13-18 kya during a period of glacial retreat in the northern Great Plains and substructuring largely occurred ~5-11 kya during a period of extreme drought cycles. A range-wide species distribution model (SDM) for R. pipiens was developed and applied to prehistoric climate conditions during the Last Glacial Maximum (21 kya) and the mid-Holocene (6 kya) from the CCSM4 climate model to identify potential refugia. The SDM indicated potential refugia existed in South Dakota or further south in Nebraska. The ancestral populations of R. pipiens in North Dakota may have inhabited these refugia, but more sampling outside the state is needed to reconstruct the route of colonization. Using microsatellite genotype data, this study determined that colonization from glacial refugia, drought dynamics in the northern Great Plains, and major rivers acting as barriers to gene flow were the defining forces shaping the regional population structure of R. pipiens in North Dakota.