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At least 199 records · Page 11Linked to original sources

A review of Arctomecon californica (Papaveraceae) with a focus on the species’ potential for propagation and reintroduction and conservation needs

Las Vegas bearpoppy ( Arctomecon californica ) occurrences have fluctuated during the past several decades, in part due to interannual variability in rainfall that influences recruitment and mortality events; yet, development in the Las Vegas Valley continues to threaten habitat supporting this species. Arctomecon californica was petitioned for listing under the Endangered Species Act in 2019 and is currently under review to determine whether listing is warranted ( USFWS 2020 ). This review updates species information for A. californica and includes recent insights into the species' seed ecology, habitat requirements and suitability models, propagation and reintroduction, and pollinator biology. We include information from the past 20 years in these areas that supplement conservation and restoration actions for the species. We also identify topics with scarce information and highlight areas for future study, including the following: preservation of genetic diversity through germplasm collections, identification of mechanisms driving the species' soil endemism, maintenance of A. californica –pollinator relationships through understanding pollinator habitat, determination of the viable seed fraction and its longevity in the soil seed reserves, and prediction of population response to regional climate change based on demographic modeling.

Arizona, Nevada↗

Trophic structure of apex fish communities in closed versus leaky lakes of arctic Alaska

Despite low species diversity and primary production, trophic structure (e.g., top predator species, predator size) is surprisingly variable among Arctic lakes. We investigated trophic structure in lakes of arctic Alaska containing arctic char Salvelinus alpinus using stomach contents and stable isotope ratios in two geographically-close but hydrologically-distinct lake clusters to investigate how these fish may interact and compete for limited food resources. Aside from different lake connectivity patterns (‘leaky’ versus ‘closed’), differing fish communities (up to five versus only two species) between lake clusters allowed us to test trophic hypotheses including: (1) arctic char are more piscivorous, and thereby grow larger and obtain higher trophic positions, in the presence of other fish species; and, (2) between arctic char size classes, resource polymorphism is more prominent, and thereby trophic niches are narrower and overlap less, in the absence of other predators. Regardless of lake cluster, we observed little direct evidence of arctic char consuming other fishes, but char were larger (mean TL = 468 vs 264 mm) and trophic position was higher (mean TP = 4.0 vs 3.8 for large char) in lakes with other fishes. Further, char demonstrated less intraspecific overlap when other predators were present whereas niche overlap was up to 100% in closed, char only lakes. As hydrologic characteristics (e.g., lake connectivity, water temperatures) will change across the Arctic owing to climate change, our results provide insight regarding potential concomitant changes to fish interactions and increase our understanding of lake trophic structure to guide management and conservation goals.

Alaska↗

Groundwater-derived nutrient and trace element transport to a nearshore Kona coral ecosystem: Experimental mixing model results

Study region The groundwater influenced coastal waters along the arid Kona coast of the Big Island, Hawai’i. Study focus A salinity-and phase partitioning-based mixing experiment was constructed using contrasting groundwater endmembers along the arid Konacoast of the Big Island, Hawai’i and local open seawater to better understand biogeochemical and physicochemical processes that influence the fate of submarine groundwater discharge (SGD)-derived nutrients and trace elements. New Hydrological Insights for the Region Treated wastewater effluent was the main source for nutrient enrichment downstream at the Honokōhau Harbor site. Conservative mixing for some constituents, such as nitrate + nitrite, illustrate the effectiveness of physical mixing to maintain oceanic concentrations in the colloid (0.02–0.45 μm) and truly dissolved (

Hawaii↗

Used-habitat calibration plots: A new procedure for validating species distribution, resource selection, and step-selection models

“Species distribution modeling” was recently ranked as one of the top five “research fronts” in ecology and the environmental sciences by ISI's Essential Science Indicators (Renner and Warton 2013), reflecting the importance of predicting how species distributions will respond to anthropogenic change. Unfortunately, species distribution models (SDMs) often perform poorly when applied to novel environments. Compounding on this problem is the shortage of methods for evaluating SDMs (hence, we may be getting our predictions wrong and not even know it). Traditional methods for validating SDMs quantify a model's ability to classify locations as used or unused. Instead, we propose to focus on how well SDMs can predict the characteristics of used locations. This subtle shift in viewpoint leads to a more natural and informative evaluation and validation of models across the entire spectrum of SDMs. Through a series of examples, we show how simple graphical methods can help with three fundamental challenges of habitat modeling: identifying missing covariates, non-linearity, and multicollinearity. Identifying habitat characteristics that are not well-predicted by the model can provide insights into variables affecting the distribution of species, suggest appropriate model modifications, and ultimately improve the reliability and generality of conservation and management recommendations.

Ecography↗

Conservation paleobiology: Leveraging knowledge of the past to inform conservation and restoration

Humans now play a major role in altering Earth and its biota. Finding ways to ameliorate human impacts on biodiversity and to sustain and restore the ecosystem services on which we depend is a grand scientific and societal challenge. Conservation paleobiology is an emerging discipline that uses geohistorical data to meet these challenges by developing and testing models of how biota respond to environmental stressors. Here we ( a ) describe how the discipline has already provided insights about biotic responses to key environmental stressors, ( b ) outline research aimed at disentangling the effects of multiple stressors, ( c ) provide examples of deliverables for managers and policy makers, and ( d ) identify methodological advances in geohistorical analysis that will foster the next major breakthroughs in conservation outcomes. We highlight cases for which exclusive reliance on observations of living biota may lead researchers to erroneous conclusions about the nature and magnitude of biotic change, vulnerability, and resilience.

Annual Review of Earth and Planetary Sciences↗

Effects of suspended sediment on the reproductive success of the tricolor shiner, a crevice-spawning minnow

Excessive sedimentation of rivers and creeks has been linked to increasing levels of imperilment in the diverse fish fauna of the southeastern United States. In particular, benthic-spawning fishes have decreased in both numbers and range. The tricolor shiner Cyprinella trichroistia is a crevice-spawning minnow that is widespread in the eastern Mobile River drainage above the Fall Line. While they are less sensitive to perturbation than the federally threatened blue shiner C. caerulea, tricolor shiners have decreased in numbers and range during the past few decades. This experiment examined the effects of 0 (control), 100, 300, and 600 mg/L initial concentrations of suspended sediment on the reproductive success of the tricolor shiner. Increasing levels of suspended sediment caused decreasing levels of reproductive success (fewer spawns and fewer eggs laid when spawning occurred). Increasing levels of suspended sediment also delayed the onset of spawning, resulting in distinct frequency distributions of clear eggs, eyed eggs, and larvae across treatments. Mortality of eggs and larvae was not a factor in the experiment. We examined daily sediment concentration data recorded at three sites in the upper Coosa River over the April-August spawning season of 1963. By weighting our experimental results by the number of days above each sediment concentration, we estimated that 20% fewer eggs were laid than would have been laid if conditions had been like those of the controls. Albeit our model is simplistic, it provides insight into how excessive sedimentation contributes to population extirpation. Mitigation of suspended sediments is an important management action for the conservation of southern Appalachian fishes that spawn in benthic habitats.

Transactions of the American Fisheries Society↗

Fish beta diversity associated with hydrologic and anthropogenic disturbance gradients in contrasting stream flow regimes

Understanding the role of hydrologic variation in structuring aquatic communities is crucial for successful conservation and sustainable management of native freshwater biodiversity. Partitioning beta diversity into the additive components of spatial turnover and nestedness can provide insight into the forces driving variability in fish assemblages across stream flow regimes. We examined stream fish beta diversity across hydrologic and anthropogenic disturbance gradients using long-term (1916–2016) site occurrence records ( n = 17,375) encompassing 252 species. We assessed total beta diversity (Sørensen dissimilarity), spatial turnover, and nestedness of fish assemblages in contrasting stream flow regimes across a gradient of decreasing flow stability: groundwater stable ( n = 77), groundwater ( n = 67), groundwater flashy ( n = 175), perennial runoff ( n = 141), runoff flashy ( n = 255), and intermittent ( n = 63) streams. Differences in total beta diversity among the stream flow regimes were driven predominantly (>86 %) by spatial turnover (i.e. species replacement) as opposed to nestedness (i.e. species loss or gain). Total fish beta diversity and spatial turnover were highest in streams with intermediate flow stability (groundwater flashy), while more flow-stable streams (groundwater stable and groundwater) had lower turnover and higher nestedness. Species turnover was also strongly associated with seasonal variation in hydrology across all flow regimes, but these relationships were most evident for assemblages in intermittent streams. Distance-based statistical comparisons showed significant correlations between beta diversity and anthropogenic disturbance variables, including dam density, dam storage volume and water withdrawals in catchments of groundwater stable streams, while hydrologic variables were more strongly correlated with beta diversity in streams with runoff-dominated and flashy flow regimes. The high spatial turnover of species implies that fish conservation actions would benefit from watershed-focused approaches targeting multiple streams with wide spatial distribution, as opposed to simply focusing on preserving sites with the greatest number of species.

Science of the Total Envionrment↗

Developing a landscape‐scale, multi‐species, and cost‐efficient conservation strategy for imperilled aquatic species in the Upper Tennessee River Basin, USA

Strategic conservation of imperilled species faces several major challenges including uncertainty in species response to management actions, budgetary constraints that limit options, and the need to scale expected conservation benefits from local to landscape levels and from single to multiple species. A structured decision‐making process was applied to address these challenges and identify a cost‐effective conservation strategy for the Federally listed endangered and threatened aquatic species in the Upper Tennessee River Basin (UTRB). The UTRB, which encompasses a landscape of ~58 000 km 2 , primarily in western North Carolina, eastern Tennessee, and south‐western Virginia, harbours one of the most globally diverse assemblages of freshwater fishes and mussels at temperate latitudes. To develop a strategy for conservation of 12 fish species and 24 mussel species over a 20‐year period, a management strategy that would best recover these species was identified given costs and uncertainty in management effectiveness. The main insights came from a trade‐off analysis that compared alternative allocations of effort among management actions. A strategy emphasizing population management, which included propagation and translocation, performed best across a wide range of objective weightings and was robust to uncertainty in management effectiveness. Species prioritization was based on the expected conservation benefit from the best performing strategy, degree of imperilment, and species‐specific management costs. Sub‐basin prioritization was based on expected conservation benefit from the best performing strategy and feasibility of habitat management and threat abatement. Although the strategy was developed for imperilled aquatic species in the UTRB, the structured process is applicable for developing cost‐efficient strategies to conserve multiple species across a landscape under uncertain management effectiveness. The process can assist a manager with limited resources to understand which species to work on, where to conduct that work, and what work would be most beneficial for those species in those catchments.

Alabama, Georgia, Kentucky, North Carolina, Tennes↗

The blind men meet the elephant at the dam: Alternative spatial and taxonomic components reveal different insights about how low-head dams impact fish biodiversity

Dams are ubiquitous environmental impacts that threaten aquatic ecosystems. The ability to compare across research studies is essential to conserve the native biodiversity that is impacted by the millions of low‐head dams that currently fragment streams and rivers. Here, we identify a previously unaddressed obstacle that impedes this generalization. Specifically, divergent spatial and taxonomic approaches that result from different conceptualizations of the dam‐biodiversity problem can produce conflicting science‐based conclusions about the same dam impact. In this research, using the same dammed and undammed sites, we evaluated the scientific generality of different conceptualizations of the dam‐biodiversity problem. We compared two different but commonly used spatial approaches—(1) above dam–below dam vs. (2) undammed–dammed comparisons—and 11 different, commonly used taxonomic approaches (three assemblage summaries, eight guilds). Sites above the dam structure had less diverse fish assemblages than sites below dams, whereas sites below the dam structure were similar to undammed sites. Thus, spatial approach 1 detected a large dam effect and spatial approach 2 detected a small dam effect. Similarly, some taxonomic responses (species richness, diversity, abundance, and number of guilds) detected large dam effects; other responses detected small (riffle specialist guild) or no dam effects (pool generalists). In summary, our results showed that how the problem was framed altered scientific conclusions and created different dam realities. The metaphor of how individual blind men disagree about the structure of an elephant, based on examinations of different body parts, reinforces the need for a coordinated, holistic perspective on dam research. Although no single approach is adequate for all problems, identifying the form, consequences of, and relationships among different research conceptualizations will set the stage for future syntheses of dam‐biodiversity research to advance science‐based conservation.

Ecosphere↗

Biodiversity connections—‘ties that bind’

Connectivity is a foundational concept in ecology and conservation and was the organising theme for the 2022 Annual Meeting of the Southeastern Fishes Council, a professional organisation dedicated to the study and conservation of freshwater fishes native to the southeast region of the United States (US). We introduce a Special Contribution of six papers selected from presentations at that meeting that illustrate perspectives on connections created by fish migration and dispersal, evolved life histories and habitat affinities and interspecific facilitation. Although focused on streams of the southeast US, each of these topics is broadly relevant to freshwater fish conservation, particularly with respect to causes and consequences of migratory fish depletion, population fragmentation and species declines. Many other connections relevant to the ecology and conservation of freshwater fishes remain relatively unexplored but could substantively advance conservation. We highlight the potential that species evolutionary histories, that is connections through time, reconstructed using species distributions and phylogenies may improve predictions of species responses to environmental change. Identifying species interdependencies, including undiscovered interactions that support survival or reproduction, could provide insights into how species losses may cascade as aquatic communities unravel. Finally, efforts to elucidate diverse connections between people and freshwater biodiversity, particularly where fisheries are historic and streams mostly go unnoticed, may prove essential to building public support for conservation measures. A research agenda anchored on ‘biodiversity connections’ has the potential to advance ecological understanding and public engagement, elements essential to conserving freshwater fishes.

Ecology of Freshwater Fish↗

A comparative analysis of common methods to identify waterbird hotspots

1. Hotspot analysis is a commonly used method in ecology and conservation to identify areas of high biodiversity or conservation concern. However, delineating and mapping hotspots is subjective and various approaches can lead to different conclusions with regard to the classification of particular areas as hotspots, complicating long-term conservation planning and implementation efforts. 2. We present a comparative analysis of recent approaches for identifying waterbird hotspots, with the goal of developing insights about the appropriate use of these methods. We selected four commonly used measures to identify persistent areas of high use: kernel density estimation, Getis-Ord Gi*, hotspot persistence, and hotspots conditional on presence, which represent the range of quantitative hotspot estimation approaches used in waterbird analyses. We applied each of the methods to aerial survey waterbird count data collected in the Great Lakes from 2012-2014 using a 5 km2 grid. For each approach, we identified areas of high use for seven species/species groups and then compared the results across all methods. 3. Our results indicate that formal hotspot analysis frameworks do not always lead to the same conclusions. The kernel density and Getis-Ord Gi* methods yielded the most similar results across all species analyzed. We found that these two models can differ substantially from the hotspot persistence and hotspots conditional on presence estimation approaches, which were not consistently similar to one another. The hotspot persistence approach differed most significantly from the other methods but is the only method to explicitly account for temporal variation. 4. We recommend considering the ecological question and scale of any conservation or management activities prior to designing survey methodologies. Deciding the appropriate definition and scale for analysis is critical for interpretation of hotspot analysis results. Combining methods using an integrative approach, either within a single analysis or post-hoc, could lead to greater consistency in the identification of waterbird hotspots.

Lake Erie, Lake Huron, Lake Michigan↗

A guide to calculating habitat-quality metrics to inform conservation of highly mobile species

Many metrics exist for quantifying the relative value of habitats and pathways used by highly mobile species. Properly selecting and applying such metrics requires substantial background in mathematics and understanding the relevant management arena. To address this multidimensional challenge, we demonstrate and compare three measurements of habitat quality: graph-, occupancy-, and demographic-based metrics. Each metric provides insights into system dynamics, at the expense of increasing amounts and complexity of data and models. Our descriptions and comparisons of diverse habitat-quality metrics provide means for practitioners to overcome the modeling challenges associated with management or conservation of such highly mobile species. Whereas previous guidance for applying habitat-quality metrics has been scattered in diversified tracks of literature, we have brought this information together into an approachable format including accessible descriptions and a modeling case study for a typical example that conservation professionals can adapt for their own decision contexts and focal populations. Considerations for Resource Managers Management objectives, proposed actions, data availability and quality, and model assumptions are all relevant considerations when applying and interpreting habitat-quality metrics. Graph-based metrics answer questions related to habitat centrality and connectivity, are suitable for populations with any movement pattern, quantify basic spatial and temporal patterns of occupancy and movement, and require the least data. Occupancy-based metrics answer questions about likelihood of persistence or colonization, are suitable for populations that undergo localized extinctions, quantify spatial and temporal patterns of occupancy and movement, and require a moderate amount of data. Demographic-based metrics answer questions about relative or absolute population size, are suitable for populations with any movement pattern, quantify demographic processes and population dynamics, and require the most data. More real-world examples applying occupancy-based, agent-based, and continuous-based metrics to seasonally migratory species are needed to better understand challenges and opportunities for applying these metrics more broadly.

Natural Resource Modeling↗

Extirpation and recolonization in a metapopulation of an endangered fish, the tidewater goby

The tidewater goby (Eucyclogobius newberryi), an endangered species in the United States, occurs in a series of isolated coastal wetlands in California. Using historical presence-absence data and our own surveys, we estimated annual rates of extirpation and recolonization for several populations of the goby in southern California. As predicted, large wetlands had lower rates of extirpation than small wetlands. There was a negative but statistically nonsignificant correlation between recolonization rate and distance to the nearest northerly source population. Populations at small sites were sensitive to drought, presumably because droughts can eliminate suitable habitat at small wetlands. Populations in small wetlands have declined over time, even after accounting for variation in stream flow, supporting the species' endangered status. Our study emphasizes the need to understand metapopulation dynamics for conserving species where the unit of conservation is a local population. It is also emphasizes the importance of not treating metapopulations as identical units. Finally, our results provide a means for describing the decline of a species that is complex in time and space and provide insight into how to target protection measures among metapopulations.

Conservation Biology↗

Evaluating contributions of recent tracking-based animal movement ecology to conservation management

The use of animal-born sensors for location-based tracking and bio-logging in terrestrial systems has expanded dramatically in the past 10 years. This rapid expansion has generated new data on how animals interact with and respond to variation in their environment, resulting in important ecological, physiological, and evolutionary insights. Although understanding the finer details of animal locations has important management relevance, applied studies are not prominent in the movement ecology literature. This is despite the long history of applied studies of animal movement and the urgent and growing need for evidence-based conservation guidance, especially in the challenging field of human-wildlife interactions. The goal of this review is to evaluate the realized contribution of tracking-based animal movement ecology to solving specific conservation problems, and to identify barriers that may hinder expansion of that contribution. To do this, we (a) briefly review the history and technologies used in animal tracking and bio-logging, (b) use a series of literature searches to evaluate the frequency with which movement ecology studies are designed to solve specific conservation problems, and (c) use this information to identify challenges that may limit the applied relevance of the field of movement ecology, and to propose pathways to expand that applied relevance. Our literature review quantifies the limited extent to which research in the field of movement ecology is designed to solve specific conservation problems, but also the fact that such studies are slowly becoming more prevalent. We discuss how barriers that limit application of these principles are likely due to constraints imposed by the types of data used commonly in the field. Problems of scale mismatch, error compounding, and data paucity all create challenges that are relevant to the field of movement ecology but may be especially pertinent in applied situations. Finding solutions to these problems will create new opportunity for movement ecologists to contribute to conservation science.

Frontiers in Ecology and Evolution↗

Application of genetics and genomics to wildlife epidemiology

Wildlife diseases can have significant impacts on wildlife conservation and management. Many of the pathogens that affect wildlife also have important implications for domestic animal and human health. However, management interventions to prevent or control wildlife disease are hampered by uncertainties about the complex interactions between pathogens and free-ranging wildlife. We often lack crucial knowledge about host ecology, pathogen characteristics, and host–pathogen dynamics. The purpose of this review is to familiarize wildlife biologists and managers with the application of genetic and genomic methodologies for investigating pathogen and host biology to better understand and manage wildlife diseases. The genesis of this review was a symposium at the 2013 annual Wildlife Society Conference. We reviewed the scientific literature and used our personal experiences to identify studies that illustrate the application of genetic and genomic methods to advance our understanding of wildlife epidemiology, focusing on recent research, new techniques, and innovative approaches. Using examples from a variety of pathogen types and a broad array of vertebrate taxa, we describe how genetics and genomics can provide tools to detect and characterize pathogens, uncover routes of disease transmission and spread, shed light on the ways that disease susceptibility is influenced by both host and pathogen attributes, and elucidate the impacts of disease on wildlife populations. Genetic and increasingly genomic methodologies will continue to contribute important insights into pathogen and host biology that will aid efforts to assess and mitigate the impacts of wildlife diseases on global health and conservation of biodiversity.

Journal of Wildlife Management↗

The evolutionary history of steelhead ( Oncorhynchus mykiss ) along the US Pacific Coast: Developing a conservation strategy using genetic diversity

Changes in genetic variation across a species range may indicate patterns of population structure resulting from past ecological and demographic events that are otherwise difficult to infer and thus provide insight into evolutionary development. Genetic data is used, drawn from 11 microsatellite loci amplified from anadromous steelhead ( Oncorhynchus mykiss ) sampled throughout its range in the eastern Pacific Ocean, to explore population structure at the southern edge in California. Steelhead populations in this region represent less than 10% of their reported historic abundance and survive in very small populations found in fragmented habitats. Genetic data derived from three independent molecular systems (allozymes, mtDNA, and microsatellites) have shown that the southernmost populations are characterized by a relatively high genetic diversity. Two hypothetical models supporting genetic population substructure such as observed were considered: (1) range expansion with founder-flush effects and subsequent population decline; (2) a second Pleistocene radiation from the Gulf of California. Using genetic and climatic data, a second Pleistocene refugium contributing to a southern ecotone seems more feasible. These data support strong conservation measures based on genetic diversity be developed to ensure the survival of this uniquely diverse gene pool.

ICES Journal of Marine Science↗

Age and sex specific timing, frequency, and spatial distribution of horseshoe crab spawning in Delaware Bay: Insights from a large-scale radio telemetry array

To study horseshoe crab Limulus polyphemus spawning behavior and migration over a large-spatial extent (>100 km), we arrayed fixed station radio receivers throughout Delaware Bay and deployed radio transmitters and archival tags on adult horseshoe crabs prior to their spawning season. We tagged and released 160 females and 60 males in 2004 and 217 females in 2005. The array covered approximately 140 km of shoreline. Recapture rates were >70% with multi-year recaptures. We categorized adult age by carapace wear. Older females tended to spawn earlier in the season and more frequently than young females, but those tendencies were more apparent in 2004 when spawning overall occurred earlier than in 2005 when spawning was delayed possibly due to decreased water temperatures. Timing of initial spawning within a year was correlated with water temperature. After adjusting for day of first spring tide, the day of first spawning was 4 days earlier for every 1 degree (̊C) rise in mean daily water temperature in May. Seventy nine % of spawning occurred during nighttime high tides. Fifty five % of spawning occurred within 3 d of a spring tide, which was slightly higher than the 47% expected if spawning was uniformly distributed regardless of tidal cycle. Within the same spawning season, males and females were observed spawning or intertidally resting at more than one beach separated by >5 km. Between years, most (77%) did not return to spawn at the same beach. Probability of stranding was strongly age dependent for males and females with older adults experiencing higher stranding rates. Horseshoe crabs staging in the shallow waters east of the channel spawned exclusively along the eastern (NJ) shoreline, but those staging west of the channel spawned throughout the bay. Overall, several insights emerged from the use of radio telemetry, which advances our understanding of horseshoe crab ecology and will be useful in conserving the Delaware Bay horseshoe crab population and habitats.

Delaware, New Jersey↗

Reducing fertilizer-nitrogen losses from rowcrop landscapes: Insights and implications from a spatially explicit watershed model

We present conceptual and quantitative models that predict changes in fertilizer-derived nitrogen delivery from rowcrop landscapes caused by agricultural conservation efforts implemented to reduce nutrient inputs and transport and increase nutrient retention in the landscape. To evaluate the relative importance of changes in the sources, transport, and sinks of fertilizer-derived nitrogen across a region, we use the spatially explicit SPAtially Referenced Regression On Watershed attributes watershed model to map the distribution, at the small watershed scale within the Upper Mississippi-Ohio River Basin (UMORB), of: (1) fertilizer inputs; (2) nutrient attenuation during delivery of those inputs to the UMORB outlet; and (3) nitrogen export from the UMORB outlet. Comparing these spatial distributions suggests that the amount of fertilizer input and degree of nutrient attenuation are both important in determining the extent of nitrogen export. From a management perspective, this means that agricultural conservation efforts to reduce nitrogen export would benefit by: (1) expanding their focus to include activities that restore and enhance nutrient processing in these highly altered landscapes; and (2) targeting specific types of best management practices to watersheds where they will be most valuable. Doing so successfully may result in a shift in current approaches to conservation planning, outreach, and funding.

Illinois, Indiana, Iowa, Minnesota, Missouri, Ohio↗