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Science to support conservation action in a large river system: The Willamette River, Oregon, USA

Management and conservation efforts that support the recovery and protection of large rivers are daunting, reflecting the complexity of the challenge and extent of effort (in terms of policy, economic investment, and spatial extent) needed to afford measurable change. These large systems have generally experienced intensive development and regulation, compromising their capacity to respond to disturbances such as climate change or wildfire. Functionally, large river and basin management require insights gained from social, ecological, geophysical, and hydrological sciences. This multi-disciplinary perspective can unveil the integrated relationship between a river network's biotic community and seasonally variable environmental conditions that are often influenced by human activities. Large rivers and their basins are constantly changing due to anthropogenic influences and as climate modifies patterns of temperature and precipitation. Because of these factors, the state of knowledge must advance to address changing conditions. The Willamette River, in western Oregon, USA, is a prime example of a basin that has experienced significant degradation and investment in rehabilitation in recent decades. Innovative science has facilitated development of fine-scale, spatially extensive datasets and models that can generate targeted conservation and rehabilitation actions that are prioritized across the entire river network. This prioritization allows investment decisions to be driven by site-specific conditions while simultaneously considering potentials for ecological improvement. Here, we review hydrologic, geomorphic, ecologic, and social conditions in the Willamette River basin through time—including pre-settlement, river development, and contemporary periods—and offer a future vision for consideration. Currently, detailed information about fish populations and habitat, hydrologic conditions, geomorphology, water quality, and land use can be leveraged to make informed decisions about protection, rehabilitation, and development. The time is ripe for strategic management and goal development for the entire Willamette River, and these efforts can be informed by comprehensive science realized through established institutions (e.g., public agencies, non-profit watershed groups, Tribes, and universities) focused on conservation and management. The approaches to science and social-network creation that were pioneered in the Willamette River basin offer insights into the development of comprehensive conservation-based planning that could be implemented in other large river systems globally.

Oregon

Whitebark pine mortality related to white pine blister rust, mountain pine beetle outbreak, and water availability

Whitebark pine ( Pinus albicaulis ) forests in the western United States have been adversely affected by an exotic pathogen ( Cronartium ribicola , causal agent of white pine blister rust), insect outbreaks ( Dendroctonus ponderosae , mountain pine beetle), and drought. We monitored individual trees from 2004 to 2013 and characterized stand-level biophysical conditions through a mountain pine beetle epidemic in the Greater Yellowstone Ecosystem. Specifically, we investigated associations between tree-level variables (duration and location of white pine blister rust infection, presence of mountain pine beetle, tree size, and potential interactions) with observations of individual whitebark pine tree mortality. Climate summaries indicated that cumulative growing degree days in years 2006–2008 likely contributed to a regionwide outbreak of mountain pine beetle prior to the observed peak in whitebark mortality in 2009. We show that larger whitebark pine trees were preferentially attacked and killed by mountain pine beetle and resulted in a regionwide shift to smaller size class trees. In addition, we found evidence that smaller size class trees with white pine blister rust infection experienced higher mortality than larger trees. This latter finding suggests that in the coming decades white pine blister rust may become the most probable cause of whitebark pine mortality. Our findings offered no evidence of an interactive effect of mountain pine beetle and white pine blister rust infection on whitebark pine mortality in the Greater Yellowstone Ecosystem. Interestingly, the probability of mortality was lower for larger trees attacked by mountain pine beetle in stands with higher evapotranspiration. Because evapotranspiration varies with climate and topoedaphic conditions across the region, we discuss the potential to use this improved understanding of biophysical influences on mortality to identify microrefugia that might contribute to successful whitebark pine conservation efforts. Using tree-level observations, the National Park Service-led Greater Yellowstone Interagency Whitebark Pine Long-term Monitoring Program provided important ecological insight on the size-dependent effects of white pine blister rust, mountain pine beetle, and water availability on whitebark pine mortality. This ongoing monitoring campaign will continue to offer observations that advance conservation in the Greater Yellowstone Ecosystem.

Ecosphere

Model-based clustering reveals patterns in central place use of a marine top predator

Satellite telemetry data are commonly used to quantify habitat selection, examine animal movements, and delineate home ranges. These data also contain valuable information concerning dens, nests, roosts, and other central places that are often associated with important life history events and may exhibit unique characteristics; however, using satellite telemetry data to study central places is complicated by common nuances like locational error and animal movement. We coupled a novel modeling framework that accounts for these nuances with an Argos satellite telemetry dataset to examine the spatiotemporal behavior associated with harbor seal haul-out sites on Kodiak Island, Alaska, USA. The methodology incorporates an observation model that accommodates multiple sources of uncertainty in telemetry data and a flexible Bayesian nonparametric model to uncover latent clustering in the telemetry locations. We also contribute extensions to examine the effect of covariates on site selection and to obtain population-level inference concerning central place use. Harbor seal haul-out sites generally occurred in inlets and bays, areas that are isolated from the open water of the Gulf of Alaska. Most individuals selected haul-out sites that were protected from wave exposure. The effects of bathymetry and shoreline complexity on haul-out site selection were variable among individual seals, as were the effects of time of day, time since low tide, and day of year on temporal patterns of haul-out use. As repositories of satellite telemetry data on a wide variety of species accumulate, so do opportunities for using this information to learn about the locations of central places, as well as the temporal patterns in their use. The model-based approach we describe offers a practical and rigorous means for gaining insight concerning these sensitive locations, knowledge of which is important for the effective management and conservation of many species.

Alaska

Comparison of microscopy and metabarcoding to identify pollen used by the critically endangered rusty patched bumble bee, Bombus affinis

Taxonomic analysis of pollen collected by bees can provide insights into their host plant use, providing information about the plant species selected for targeted conservation strategies. The two main identification approaches used are morphological analysis of pollen samples affixed to microscope slides (i.e., microscopic palynology) and molecular analysis of samples. Both methods are widely used for freshly collected materials and have been compared in multiple studies, yet their application to archived samples remains to be explored. Archived samples may be particularly useful for the study of rare or protected species, particularly when historical foraging patterns are uncertain. We used both methods to analyse pollen collected by the endangered rusty patched bumble bee species, Bombus affinis Cresson, applied to museum-archived specimens. Pollen samples were removed from the corbiculae of bumble bees originally collected in Michigan between 1914 and 1974. Samples included 24 rusty patched bumble bees each with large pollen loads on both corbiculae, allowing for pollen from the same bee to be analysed using both methods. DNA metabarcoding detected more plant taxa than light microscopy, and DNA barcoding also had higher taxonomic resolution when compared to taxa determined using light microscopy. In many instances, pollen could only be confidently identified to tribe or family with light microscopy. Discrepancy between methods decreased when taxa identified via DNA metabarcoding were binned into ecologically relevant groups corresponding to those identified using light microscopy. Although binning demonstrated smaller within-method variance, there was still minimal correspondence between the two methods. Results indicate there are benefits and biases unique to each method and highlight the utility of binning taxonomic results to morphological or ecological groupings.

Insect Conservation and Diversity

Characteristics of foraging sites and protein status in wintering muskoxen: insights from isotopes of nitrogen

Identifying links between nutritional condition of individuals and population trajectories greatly enhances our understanding of the ecology, conservation, and management of wildlife. For northern ungulates, the potential impacts of a changing climate to populations are predicted to be nutritionally mediated through an increase in the severity and variance in winter conditions. Foraging conditions and the availability of body protein as a store for reproduction in late winter may constrain productivity in northern ungulates, yet the link between characteristics of wintering habitats and protein status has not been established for a wild ungulate. We used a non‐invasive proxy of protein status derived from isotopes of N in excreta to evaluate the influence of winter habitats on the protein status of muskoxen in three populations in Alaska (2005–2008). Multiple regression and an information‐theoretic approach were used to compare models that evaluated the influence of population, year, and characteristics of foraging sites (components of diet and physiography) on protein status for groups of muskoxen. The observed variance in protein status among groups of muskoxen across populations and years was partially explained (45%) by local foraging conditions that affected forage availability. Protein status improved for groups of muskoxen as the amount of graminoids in the diet increased (−0.430 ± 0.31, β± 95% CI) and elevation of foraging sites decreased (0.824 ± 0.67). Resources available for reproduction in muskoxen are highly dependent upon demographic, environmental, and physiographic constraints that affect forage availability in winter. Due to their very sedentary nature in winter, muskoxen are highly susceptible to localized foraging conditions; therefore, the spatial variance in resource availability may exert a strong effect on productivity. Consequently, there is a clear need to account for climate–topography effects in winter at multiple scales when predicting the potential impacts of climatic shifts on population trajectories of muskoxen.

Oikos

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

Salmon data mobilization

Despite substantial research and conservation efforts, many salmon populations are in decline. Globally, salmon research is not delivering effective decision support products to help managers apply research insights as informed management actions. Data Mobilization (DM) is a key step towards building the wider evidence base required to deliver accountable, reliable, and usable scientific advice to managers. Best practices for DM are being adopted throughout the scientific community but have not permeated deeply into the culture of salmon research and conservation. To address this, we present a strategy for Salmon Data Mobilization (SDM). This strategy defines three spheres of agencies and practitioners that must interact to advance SDM: (1) authoritative bodies that can create policies to support SDM uptake; (2) agencies that can promote, fund, and implement those policies; and (3) the broad salmon community of practice that can support uptake of SDM within focused interest groups. We sketch a future for SDM and propose functional changes required to improve it throughout the community.

North Pacific Anadromous Fish Commission Bulletin

Using a genetic mixture model to study phenotypic traits: Differential fecundity among Yukon river Chinook Salmon

Fecundity is a vital population characteristic that is directly linked to the productivity of fish populations. Historic data from Yukon River (Alaska) Chinook salmon Oncorhynchus tshawytscha suggest that length‐adjusted fecundity differs among populations within the drainage and either is temporally variable or has declined. Yukon River Chinook salmon have been harvested in large‐mesh gill‐net fisheries for decades, and a decline in fecundity was considered a potential evolutionary response to size‐selective exploitation. The implications for fishery conservation and management led us to further investigate the fecundity of Yukon River Chinook salmon populations. Matched observations of fecundity, length, and genotype were collected from a sample of adult females captured from the multipopulation spawning migration near the mouth of the Yukon River in 2008. These data were modeled by using a new mixture model, which was developed by extending the conditional maximum likelihood mixture model that is commonly used to estimate the composition of multipopulation mixtures based on genetic data. The new model facilitates maximum likelihood estimation of stock‐specific fecundity parameters without first using individual assignment to a putative population of origin, thus avoiding potential biases caused by assignment error. The hypothesis that fecundity of Chinook salmon has declined was not supported; this result implies that fecundity exhibits high interannual variability. However, length‐adjusted fecundity estimates decreased as migratory distance increased, and fecundity was more strongly dependent on fish size for populations spawning in the middle and upper portions of the drainage. These findings provide insights into potential constraints on reproductive investment imposed by long migrations and warrant consideration in fisheries management and conservation. The new mixture model extends the utility of genetic markers to new applications and can be easily adapted to study any observable trait or condition that may vary among populations.

Transactions of the American Fisheries Society

Wyoming Landscape Conservation Initiative—A case study in partnership development

The Wyoming Landscape Conservation Initiative (WLCI) is a successful example of collaboration between science and natural resource management at the landscape scale. In southwestern Wyoming, expanding energy and mineral development, urban growth, and other changes in land use over recent decades, combined with landscape-scale drivers such as climate change and invasive species, have presented compelling challenges to resource managers and a diverse group of Federal, State, industry, and non-governmental organizations, as well as citizen stakeholders. To address these challenges, the WLCI was established as a collaborative forum and interagency partnership to develop and implement science-based conservation actions. About a decade after being established, this report documents the establishment and history of the WLCI, focusing on the path to success of the initiative and providing insights and details that may be useful in developing similar partnerships in other locations. Not merely retrospective, the elements of the WLCI that are presented herein are still in play, still evolving, and still contributing to the resolution of compelling conservation challenges in the Western United States. The U.S. Geological Survey has developed many successful longstanding partnerships, of which the WLCI is one example. “As the Nation’s largest water, earth, and biological science and civilian mapping agency, the U.S. Geological Survey collects, monitors, analyzes, and provides scientific understanding about natural resource conditions, issues, and problems. The diversity of our scientific expertise enables us to carry out large-scale, multi-disciplinary investigations and provide impartial scientific information to resource managers, planners, and other customers” (U.S. Geological Survey, 2016).

Wyoming

Pit tag application in native freshwater mussels: Case studies across small, medium, and large rivers

Since their first use in the mid-1980s, external passive integrated transponder (PIT) tags have facilitated innovative investigations into multiple biological traits of animals. For native freshwater mussels, PIT tags are frequently used in capture-mark-recapture applications because they allow repeated, noninvasive sampling, are easy to apply, have high retention rates, and have negligible short-term effects on growth and survival. Because of these traits, resource managers and scientists are using PIT-tagged animals to estimate survival and movement of mussels associated with restoration efforts. However, consistency is limited in how PIT tags are affixed, monitored, and reported. Thus, our objectives were to (1) share our collective experiences in PIT tagging mussels across three case studies in small, medium, and large rivers and (2) propose guidelines for tagging and reporting data from PIT tag studies with native freshwater mussels to facilitate comparisons across future studies. The number of studies that have marked mussels with PIT tags has increased over the past 10 years. The ability to detect mussels using PIT tags has substantially advanced research in three areas of mussel ecology: (1) estimating vital rates (e.g., growth and survival), (2) tracking movements and behaviors of captively propagated, wild, and translocated individuals, and (3) improving our understanding of life history traits, such as reproductive timing. Each case study offers insights on tagging methods, tag loss, tag retention, and monitoring frequency across multiple species that range in conservation status from common to rare. We conclude with best-practice guidelines for placing PIT tags on freshwater mussels and a list of variables that could be reported in future studies to facilitate cross-system comparisons.

Freshwater Mollusk Biology and Conservation

Comparison of beaver density estimates from aerial surveys of waterways versus transects

Historic beaver-sign (Castor canadensis) survey flights were often conducted over waterways to maximize beaver detections. However, densities determined from strip transect surveys are more useful to compare across and within study areas than waterway indices based on observations per distance flown because transects are more representative of the wider landscape. Yet, it is unknown if, and to what extent, aerial waterway surveys are reflective of transect densities. I conducted aerial surveys for active beaver sign each fall during 2015–2018 over two waterway routes and two corresponding strip-transect routes. The simple linear regression of transect densities on waterway densities (n=8) yielded a reasonable (R2=0.79) preliminary equation for converting historic waterway data to transect densities. Additionally, visual inspection indicated that converted waterway densities reasonably reflected the trend in transect densities in an area where the wider habitat was similar in terms of beaver harvest, land use, and proportion of water features. Although trend was well-reflected, individual waterway densities in this area were only 57-75% of transect densities. In other areas, where water features were limited, visual inspection suggested the trend of waterway densities was less reflective of transect density trend and individual waterway densities overestimated transect densities (up to 309%). Nevertheless, while transect densities are better for comparisons within and across study areas, waterway surveys are still important for timely and specific within-study area insights. This research provides useful benchmark examples of reliability regarding waterway observation indices converted to densities for conservation, research, and management of beaver and their ecosystems. Because these conclusions are based on a small sample, additional research is recommended to better define this relationship especially in areas with differing habitat, beaver harvest, land use patterns, etc.

Minnesota

The transformative impact of genomics on sage-grouse conservation and management

For over two decades, genetic studies have been used to assist in the conservation and management of both Greater Sage-grouse ( Centrocercus urophasianus ) and Gunnison Sage-grouse ( C. minimus ), addressing a wide variety of topics including taxonomy, parentage, population connectivity, and demography. The field of conservation genetics has been transformed by dramatic improvements in sequencing technology, facilitating genomic studies in many wildlife species. The quality and amount of data generated by genomic methods vastly exceed that of traditional genetic studies, allowing for increased precision in estimating genetic parameters of interest. Perhaps more importantly, genomic methods can provide insight into non-neutral evolution such as adaptive divergence. Here we recount the shift from genetic to genomic methods using two wildlife species of substantial conservation interest, focusing on the improved capabilities and advantages of genomic methods. For instance, reassessment of divergence in sage-grouse using genomic methods confirmed strong differentiation between the two species and revealed that a small population in the state of Washington was more genetically distinct than previously recognized. Further, new genomic resources and approaches have been used to identify a family of genes linked to local dietary adaptation suggesting that sage-grouse may possess digestive and metabolic adaptations that mitigate the effects of consuming plant secondary metabolites like those found in sagebrush. Genetic variation among populations in these gene regions is thought to be involved with local dietary adaptations, and therefore maintaining the tie between sage-grouse and the chemistry of local sagebrush may be an important management consideration. We posit that the integration of newly developed genomic resources combined with the vast wealth of ecological and behavioral data for sage-grouse has the potential to shed light on mechanistic relationships that ultimately are vital to the conservation and management of these species.

Book chapter

Wildlife value orientation of landowners from five states in the upper midwest, USA

Five Upper Midwest states (Montana, North Dakota, South Dakota, Minnesota, Iowa) participated in a Plains and Prairie Landscape Conservation Cooperative (PPP-LCC) funded survey of landowners. All five surveys included a 13-item wildlife value orientation (WVO) scale to provide insight into how landowners in this region make land use decisions that affect wildlife habitat. Most landowners were utilitarian (59%), 11% were identified as mutualists, and pluralists and distanced landowners were 15% each. Pluralist and mutualist landowners placed more importance on wildlife considerations when making land use decisions compared to utilitarian and distanced landowners. However, landowners’ WVO was not related to their participation in United States Department of Agriculture (USDA) Farm Bill conservation programs. The proportion of WVO types among landowners varied considerably by age cohort. The proportion of utilitarians was highest for landowners born during the 1970s age cohorts and has since declined.

Iowa, Minnesota, Montana, North Dakota, South Dako

Spatially-structured statistical network models for landscape genetics

A basic understanding of how the landscape impedes, or creates resistance to, the dispersal of organisms and hence gene flow is paramount for successful conservation science and management. Spatially structured ecological networks are often used to represent spatial landscape‐genetic relationships, where nodes represent individuals or populations and resistance to movement is represented using non‐binary edge weights. Weights are typically assigned or estimated by the user, rather than observed, and validating such weights is challenging. We provide a synthesis of current methods used to estimate edge weights and an overview of common model types, stressing the advantages and disadvantages of each approach and their ability to model landscape‐genetic data. We further explore a set of spatial‐statistical methods that provide ecologists with alternative approaches for modeling spatially explicit processes that may affect genetic structure. This includes an overview of spatial autoregressive models, with a particular focus on how correlation and partial correlation are used to represent neighborhood structure with the inverse of the covariance matrix (i.e., precision matrix). We then demonstrate how to model resistance by specifying an appropriate statistical model on the nodes, conditioned on the edge weights, through the precision matrix. This integration of network ecology and spatial statistics provides a practical analytical framework for landscape‐genetic studies. The results can be used to make statistical inferences about the relative importance of individual landscape characteristics, such as the vegetative cover, hillslope, or the presence of roads or rivers, on gene flow. In addition, the R code we include allows readers to explore landscape‐genetic structure in their own datasets, which will potentially provide new insights into the evolutionary processes that generated ecological networks, as well as valuable information about the optimal characteristics of conservation corridors.

Ecological Monographs

Making decisions in complex landscapes: Headwater stream management across multiple federal agencies

Headwater stream ecosystems are vulnerable to numerous threats associated with climate and land use change. In the northeastern US, many headwater stream species (e.g., brook trout and stream salamanders) are of special conservation concern and may be vulnerable to climate change influences, such as changes in stream temperature and streamflow. Federal land management agencies (e.g., US Fish and Wildlife Service, National Park Service, USDA Forest Service, Bureau of Land Management and Department of Defense) are required to adopt policies that respond to climate change and may have longer-term institutional support to enforce such policies compared to state, local, non-governmental, or private land managers. However, federal agencies largely make management decisions in regards to headwater stream ecosystems independently. This fragmentation of management resources and responsibilities across the landscape may significantly impede the efficiency and effectiveness of conservation actions, and higher degrees of collaboration may be required to achieve conservation goals. This project seeks to provide an example of cooperative landscape decision-making to address the conservation of headwater stream ecosystems. We identified shared and contrasting objectives of each federal agency and potential collaboration opportunities that may increase efficient and effective management of headwater stream ecosystems in two northeastern US watersheds. These workshops provided useful insights into the adaptive capacity of federal institutions to address threats to headwater stream ecosystems. Our ultimate goal is to provide a decision-making framework and analysis that addresses large-scale conservation threats across multiple stakeholders, as a demonstration of cooperative landscape conservation for aquatic ecosystems. Additionally, we aim to provide new scientific knowledge and a regional perspective to resource managers to help inform local management decisions.

Maine, New Hampshire, Virginia, West Virginia

Early season tropical cyclones affect birds breeding on a barrier island

Animal populations often experience acute natural disturbances, most of which are connected to short—term weather events. Occurrences of early—season tropical cyclones during the peak of the avian breeding season are likely to increase with climate change, which can substantially impact populations of coastal breeding birds at multiple scales. To understand the acute impacts of severe tropical cyclones, we investigated how abundances of breeding birds changed before and after 4 early season named tropical cyclones on a barrier island in the Gulf of Mexico. We detected a change in pre— versus post—storm numbers of breeding birds after 2 out of 4 storms between 2015–2020. Following Tropical Storm Cindy and Hurricane Barry, significant declines in breeding adults ranged from a reduction of 32 Willets ( Tringa semipalmata ) to 105 Wilson’s Plover ( Anarhynchus wilsoni a) and 18 Willets to 1,794 Black Skimmer ( Rynchops niger ), respectively. The lack of response following 2 storms was likely due to the location of the storms relative to the island. Following the 2 storms that did elicit a response by breeding birds, we observed lower abundances in the majority of species after the storm passed. Species life histories and habitat restoration might explain the species—specific responses we observed. Our study documents occurrences of early—season tropical cyclones negatively affecting coastal breeding birds, which could be exacerbated with sea level rise. Additionally, our findings may provide insights regarding island design and nest susceptibility to flooding events, which may aid land managers as well as conservation and restoration planners.

Louisiana

Quality of groundwater in the Denver Basin aquifer system, Colorado, 2003-5

Groundwater resources from alluvial and bedrock aquifers of the Denver Basin are critical for municipal, domestic, and agricultural uses in Colorado along the eastern front of the Rocky Mountains. Rapid and widespread urban development, primarily along the western boundary of the Denver Basin, has approximately doubled the population since about 1970, and much of the population depends on groundwater for water supply. As part of the National Water-Quality Assessment Program, the U.S. Geological Survey conducted groundwater-quality studies during 2003–5 in the Denver Basin aquifer system to characterize water quality of shallow groundwater at the water table and of the bedrock aquifers, which are important drinking-water resources. For the Denver Basin, water-quality constituents of concern for human health or because they might otherwise limit use of water include total dissolved solids, fluoride, sulfate, nitrate, iron, manganese, selenium, radon, uranium, arsenic, pesticides, and volatile organic compounds. For the water-table studies, two monitoring-well networks were installed and sampled beneath agricultural (31 wells) and urban (29 wells) land uses at or just below the water table in either alluvial material or near-surface bedrock. For the bedrock-aquifer studies, domestic- and municipal-supply wells completed in the bedrock aquifers were sampled. The bedrock aquifers, stratigraphically from youngest (shallowest) to oldest (deepest), are the Dawson, Denver, Arapahoe, and Laramie-Fox Hills aquifers. The extensive dataset collected from wells completed in the bedrock aquifers (79 samples) provides the opportunity to evaluate factors and processes affecting water quality and to establish a baseline that can be used to characterize future changes in groundwater quality. Groundwater samples were analyzed for inorganic, organic, isotopic, and age-dating constituents and tracers. This report discusses spatial and statistical distributions of chemical constituents and evaluates natural and human-related processes that affect water quality. Findings are synthesized to assess the vulnerability of the Denver Basin aquifer system to groundwater contamination. The chemistry of groundwater samples collected from the water-table wells was generally different from that of samples collected from the bedrock-aquifer wells. Samples from the water-table wells tended to have higher concentrations of total dissolved solids and most major ions. Concentrations of several constituents with potential human-health concerns, including nitrate, selenium, uranium, and arsenic, decreased with depth and were highest in samples from the water-table wells. Exceedances of drinking-water standards and water-quality benchmarks were more frequently associated with shallow groundwater samples; concentrations of total dissolved solids and sulfate exceeded water-quality benchmarks for about half or more of samples from the water-table wells. The sediments and rocks of the Denver Basin are natural sources of the trace elements selenium, uranium, and arsenic, which affect their concentrations in groundwater. Detections of organic contaminants, which are typically indicative of human sources of contamination to groundwater, were more frequent in samples from the water-table wells. Pesticide compounds and volatile organic compounds were detected in 33 and 62 percent, respectively, of water-table well samples. Detected organic contaminant concentrations were much less than the associated drinking-water standards. Samples collected from the bedrock aquifers had lower concentrations of total dissolved solids than did samples collected from the water-table wells, although within the bedrock-aquifer samples, concentrations increased from the Dawson to Denver to Arapahoe to Laramie-Fox Hills aquifers. Concentrations of total dissolved solids and many constituents varied spatially and with depth in the bedrock aquifers, likely as a result of ion-exchange and oxidation-reduction reactions, which are important processes affecting water quality. Major-ion chemistry generally evolved from a calcium-bicarbonate to calcium-sulfate composition, with some sodium-bicarbonate and sodium-sulfate facies in the deeper bedrock aquifers, likely resulting from longer residence times and more extensive water-rock interaction. Oxidation-reduction conditions generally evolved from oxic at the water table to anoxic with increasing depth in the bedrock aquifers. Most samples from the bedrock aquifers were anoxic. Exceedances of drinking-water standards and water-quality benchmarks for the bedrock aquifers occurred in 1 percent or less of samples for nitrate, selenium, or arsenic; there were no exceedances for uranium. Exceedances for total dissolved solids, sulfate, manganese, and iron were generally between about 10 and 20 percent for the bedrock-aquifer samples. Radon concentrations, which were only measured in samples collected from two of the bedrock aquifers, exceeded the lower proposed drinking-water standard for more than 90 percent of samples but exceeded the higher alternative standard for less than 5 percent of samples. Pesticide compounds and volatile organic compounds were detected in 3 and 22 percent, respectively, of bedrock-aquifer samples, all at concentrations that were that were much less than drinking-water standards. Water-quality data were synthesized to evaluate factors that affect spatial and depth variability in water quality and to assess aquifer vulnerability to contaminants from geologic materials and those of human origin. The quality of shallow groundwater in the alluvial aquifer and shallow bedrock aquifer system has been adversely affected by development of agricultural and urban areas. Land use has altered the pattern and composition of recharge. Increased recharge from irrigation water has mobilized dissolved constituents and increased concentrations in the shallow groundwater. Concentrations of most constituents associated with poor or degraded water quality in shallow groundwater decreased with depth; many of these constituents are not geochemically conservative and are affected by geochemical reactions such as oxidation-reduction reactions. Groundwater age tracers provide additional insight into aquifer vulnerability and help determine if young groundwater of potentially poor quality has migrated to deeper parts of the bedrock aquifers used for drinking-water supply. Age-tracer results were used to group samples into categories of young, mixed, and old groundwater. Groundwater ages transitioned from mostly young in the water-table wells to mostly mixed in the shallowest bedrock aquifer, the Dawson aquifer, to mostly old in the deeper bedrock aquifers. Although the bedrock aquifers are mostly old groundwater of good water quality, several lines of evidence indicate that young, contaminant-bearing recharge has reached shallow to moderate depths in some areas of the bedrock aquifers. The Dawson aquifer is the most vulnerable of the bedrock aquifers to contamination, but results indicate that the older (deeper) bedrock aquifers are also vulnerable to groundwater contamination and that mixing with young recharge has occurred in some areas. Heavy pumping has caused water-level declines in the bedrock aquifers in some parts of the Denver Basin, which has the potential to enhance the transport of contaminants from overlying units. Results of this study are consistent with the existing conceptual understanding of aquifer processes and groundwater issues in the Denver Basin and add new insight into the vulnerability of the bedrock aquifers to groundwater contamination.

Colorado

Disease-smart climate adaptation for wildlife management and conservation

Climate change is a well-documented driver and threat multiplier of infectious disease in wildlife populations. However, wildlife disease management and climate-change adaptation have largely operated in isolation. To improve conservation outcomes, we consider the role of climate adaptation in initiating or exacerbating the transmission and spread of wildlife disease and the deleterious effects thereof, as illustrated through several case studies. We offer insights into best practices for disease-smart adaptation, including a checklist of key factors for assessing disease risks early in the climate adaptation process. By assessing risk, incorporating uncertainty, planning for change, and monitoring outcomes, natural resource managers and conservation practitioners can better prepare for and respond to wildlife disease threats in a changing climate.

Frontiers in Ecology and the Environment