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Variation and plasticity and their interaction with urbanization in Guadalupe Bass populations on and off the Edwards Plateau

The Colorado River Basin in Texas has experienced major alterations to its hydrologic regime due to changing land and water use patterns. These anthropogenic influences on hydrologic variability have had major implications for riparian and aquatic ecosystems and the species dependent upon them. However, impacts are often assessed at a limited temporal and spatial scale, tending to focus on relatively short and discrete periods or portions of a river basin. It is not clear how basin-wide alterations occurring over decades affect species. Guadalupe Bass Micropterus treculii are endemic to central Texas and are typically associated with shallow runs and riffles in small streams. However, Guadalupe Bass are found throughout the Colorado River Basin, including the mainstem portion of the lower river downstream of the city of Austin where they support a popular fishery. Because Guadalupe Bass exist across a wide range of stream orders within the basin, it is unclear whether populations respond similarly to anthropogenic disturbances or to conservation and restoration activities. Therefore, our objectives were to: Assess the effects of urbanization and hydrology on the population structure and dynamics of Guadalupe Bass. Evaluate the effects of environmental gradients on ecomorphological variation in Guadalupe Bass populations across multiple spatial scales. Describe the life history, habitat use, and behavior of the Guadalupe Bass population in the lower Colorado River and compare it to populations in more “typical” habitats. Results contribute to an understanding of the response of Guadalupe Bass to anthropogenic disturbances, including increased urbanization in central Texas and further assist in the conservation of the species. The ability of the population to not only persist, but flourish downstream of a heavily populated urban area presented a unique opportunity to investigate a native species response to anthropogenic disturbance. This research revealed differences in Guadalupe Bass habitat associations and movements, contrasts in age and growth, and morphological variation across a gradient of disturbance throughout the Colorado River Basin. Results of this work provide information on the potential effects of human population growth and increased water withdrawals on Guadalupe Bass populations. Additionally, this work adds to an understanding of the unique Guadalupe Bass population found in the lower Colorado River and how it differs from upstream tributary populations. Gathering additional population-level information facilitates conservation actions critical to preserving preferred habitat and promoting growth rates for Guadalupe Bass in streams of different sizes and flow conditions while highlighting interpopulation differences that may warrant consideration for stocking programs and other management strategies. Key findings of this study were: The similarity in response of growth rates to streamflow throughout the Colorado Basin suggests phenotypic plasticity in this trait rather than population-specific adaptations. Reductions in streamflows in the Colorado River Basin, whether due to increased frequency of drought or increased anthropogenic water withdrawal, will likely result in lower Guadalupe Bass growth rates with the potential to impact the structure of populations. Growth and recruitment showed a positive correlation with increased baseflows and mean monthly flows; however, continued assessment is necessary to determine a true relationship. We documented morphological divergence among Guadalupe Bass populations in response to spatial and temporal environmental variation. These ecomorphological differences among populations provide insight into the ability of Guadalupe Bass to respond to the differing in-stream habitat and flow conditions between small ‘typical’ tributary systems and the mainstem Colorado River. Morphological variation may be a population-level adaptation that potentially needs to be taken into consideration when choosing broodstock to maximize stocking success within a system. Understanding the morphological differences between Guadalupe Bass populations in response to local conditions could improve the success of restoration and supplemental stocking programs, especially in the ever-changing landscape of central Texas. We established a baseline for understanding the morphological response of Guadalupe Bass to increased population growth and the threats posed by increased water withdrawals and impervious surface. The mainstem population of Guadalupe Bass was generally more mobile, and more responsive to changes in streamflow, than tributary populations. The observed differences could influence the response of Guadalupe Bass populations to conservation and management actions, such as habitat restoration efforts. Continued monitoring of recruitment and angler exploitation may be beneficial to identify any changes that could negatively impact the population. Conservation initiatives solely focused on physical instream or riparian habitat are unlikely to be as beneficial to Guadalupe Bass as those focused on restoring or maintaining adequate streamflow

Cooperator Science Series↗

Photometric properties of Titan's surface from Cassini VIMS: Relevance to titan's hemispherical albedo dichotomy and surface stability

The Visual and Infrared Mapping Spectrometer (VIMS) instrument on the Cassini Saturn Orbiter returned spectral imaging data as the spacecraft undertook six close encounters with Titan beginning 7 July, 2004. Three of these flybys each produced overlapping coverage of two distinct regions of Titan's surface. Twenty-four points were selected on approximately opposite hemispheres to serve as photometric controls. Six points were selected in each of four reflectance classes. On one hemisphere each control point was observed at three distinct phase angles. From the derived phase coefficients, preliminary normal reflectances were derived for each reflectance class. The normal reflectance of Titan's surface units at 2.0178 ??m ranged from 0.079 to 0.185 for the most absorbing to the most reflective units assuming no contribution from absorbing haze. When a modest haze contribution of ??=0.1 is considered these numbers increase to 0.089-0.215. We find that the lowest three reflectance classes have comparable normal reflectance on either hemisphere. However, for the highest brightness class the normal reflectance is higher on the hemisphere encompassing longitude 14-65?? compared to the same high brightness class for the hemisphere encompassing 122-156?? longitude. We conclude that an albedo dichotomy observed in continental sized units on Titan is due not only to one unit having more areal coverage of reflective material than the other but the material on the brighter unit is intrinsically more reflective than the most reflective material on the other unit. This suggests that surface renewal processes are more widespread on Titan's more reflective units than on its less reflective units. We note that one of our photometric control points has increased in reflectance by 12% relative to the surrounding terrain from July of 2004 to April and May of 2005. Possible causes of this effect include atmospheric processes such as ground fog or orographic clouds; the suggestion of active volcanism cannot be ruled out. Several interesting circular features which resembled impact craters were identified on Titan's surface at the time of the initial Titan flyby in July of 2004. We traced photometric profiles through two of these candidate craters and attempted to fit these profiles to the photometric properties expected from model depressions. We find that the best-fit attempt to model these features as craters requires that they be unrealistically deep, approximately 70 km deep. We conclude that despite their appearance, these circular features are not craters, however, the possibility that they are palimpsests cannot be ruled out. We used two methods to test for the presence of vast expanses of liquids on Titan's surface that had been suggested to resemble oceans. Specular reflection of sunlight would be indicative of widespread liquids on the surface; we found no evidence of this. A large liquid body should also show uniformity in photometric profile; we found the profiles to be highly variable. The lack of specular reflection and the high photometric variability in the profiles across candidate oceans is inconsistent with the presence of vast expanses of flat-lying liquids on Titan's surface. While liquid accumulation may be present as small, sub-pixel-sized bodies, or in areas of the surface which still remain to be observed by VIMS, the presence of large ocean-sized accumulations of liquids can be ruled out. The Cassini orbital tour offers the opportunity for VIMS to image the same parts of Titan's surface repeatedly at many different illumination and observation geometries. This creates the possibility of understanding the properties of Titan's atmosphere and haze by iteratively adapting models to create a best fit to the surface reflectance properties. ?? 2006 Elsevier Ltd. All rights reserved.

Planetary and Space Science↗

Groundwater quality, age, and susceptibility and vulnerability to nitrate contamination with linkages to land use and groundwater flow, Upper Black Squirrel Creek Basin, Colorado, 2013

The Upper Black Squirrel Creek Basin is located about 25 kilometers east of Colorado Springs, Colorado. The primary aquifer is a productive section of unconsolidated deposits that overlies bedrock units of the Denver Basin and is a critical resource for local water needs, including irrigation, domestic, and commercial use. The primary aquifer also serves an important regional role by the export of water to nearby communities in the Colorado Springs area. Changes in land use and development over the last decade, which includes substantial growth of subdivisions in the Upper Black Squirrel Creek Basin, have led to uncertainty regarding the potential effects to water quality throughout the basin. In response, the U.S. Geological Survey, in cooperation with Cherokee Metropolitan District, El Paso County, Meridian Service Metropolitan District, Mountain View Electric Association, Upper Black Squirrel Creek Groundwater Management District, Woodmen Hills Metropolitan District, Colorado State Land Board, and Colorado Water Conservation Board, and the stakeholders represented in the Groundwater Quality Study Committee of El Paso County conducted an assessment of groundwater quality and groundwater age with an emphasis on characterizing nitrate in the groundwater. Groundwater-quality samples were collected from 50 randomly selected wells between May and June 2013. The samples were analyzed for major ions, nutrients, dissolved gases, tritium ( 3 H), chlorofluorocarbons (CFC-11, CFC-12, and CFC-113), and fuel products (such as benzene, toluene, ethylbenzene, and xylenes). None of the groundwater samples exceeded the U.S. Environmental Protection Agency (EPA) National Primary Drinking Water Regulations for primary maximum contaminant levels (MCL) for major ions. Secondary maximum contaminant levels, which are not health concerns and affect mainly taste, color, or odor of the water, were observed in rare instances for pH (2 samples), chloride (1 sample), iron (3 samples), and manganese (8 samples). The secondary maximum contaminant level for total dissolved solids was also exceeded for two samples. Nitrate (nitrite plus nitrate as nitrogen in groundwater) was elevated above the estimated background concentration of natural recharge waters of 1 milligram per liter (mg/L) in 44 of the 50 wells sampled and showed a median concentration of 5.4 mg/L. Nitrate concentrations were above the MCL of 10 mg/L in 5 of the 50 wells sampled and above half of the EPA MCL (5 mg/L) in 27 of the 50 wells sampled, which included samples above the MCL. Dissolved-oxygen concentrations exceeded 0.5 mg/L in 95 percent of reported values (40 of 42 samples) and exceeded 2.0 mg/L in 90 percent of reported values (38 of 42 samples). The oxidized conditions observed in most areas indicate that nitrate from fertilizers and animal or human waste was geochemically stable and could persist in the groundwater for decades or perhaps longer. A historical analysis of median nitrate concentrations over nearly three decades showed an increase in nitrate of approximately 1 mg/L from 4.3 to 5.4 mg/L, although the increase was not determined to be significantly different using nonparametric statistical methods. Major-ion data indicate that groundwater representative of the primary aquifer was classified as calcium-sodium bicarbonate type water. Other water samples from wells located mainly along the periphery of the primary aquifer had cation-anion compositions consistent with distinct water sources, including groundwater contributions from the underlying bedrock aquifers. The areas with differentiable water sources were located mainly where alluvial deposits were thin and geologic contacts to the underlying bedrock aquifers were relatively shallow. Nitrate concentrations in the groundwater were evaluated for relations to land use. An agricultural region was defined using a sequence of land satellite imagery. Groundwater flow directions interpreted from median water-table elevations measured from 2000 to 2013 were used in conjunction with cropland locations to define the agricultural region boundaries by encompassing potential pathways of nitrate transport in the groundwater from nitrogen-based fertilizers. A statistically significant higher median nitrate concentration was observed for areas inside the agricultural region (6.7 mg/L) compared to areas outside the agricultural region (2.3 mg/L), although median concentrations in both areas were below the MCL (10 mg/L). Median nitrate concentration was also significantly greater in land parcels with septic use (4.9 mg/L) compared to nonseptic parcels (1.7 mg/L). In general, agriculture or septic use was identified as the primary source of nitrate, depending on location, while commercial, county, grazing, and residential land uses were generally secondary sources of nitrate. Apparent groundwater ages were estimated from chlorofluorocarbons (CFC-11, CFC-12, and CFC-113) and tritium ( 3 H) data using models that assumed piston flow and binary mixing (dilution of a young component with old, tracer-free water). The mean and median groundwater ages were about 30 years and the standard deviation was 6 years, indicating that most groundwater in the primary aquifer was “young” water that had recharged to the aquifer over the last few decades (post-1950s). The median fraction of young water was about 71 percent, and the standard deviation was 29 percent. The remaining water predated the 1950s, which may have originated from deeper geologic formations or may represent slow moving groundwater within the primary aquifer. Some of the oldest groundwater ages (older than 30 years) were observed in the upper reaches of the aquifer to the northwest where the primary aquifer is thin and intersects bedrock, supporting the hypothesis of geochemically distinct groundwater entering the primary aquifer from below. Groundwater that had reached the central part of the aquifer from upgradient areas of the basin was variable in age because of differences in flow paths and travel velocities. The groundwater age analysis showed that current (2013) land-use practices could affect water quality over decades to come, and that responses to remedial actions could be slow, especially for constituents, such as nitrate, that are stable under oxidized conditions. Fuel products (including acetone, benzene, diisopropyl ether, ethylbenzene, methyl acetate, methyl tertiary butyl ether (MTBE), methyl tert-pentyl ether, m- + p-xylene, o-xylene, tert-amyl alcohol, tert-butyl alcohol, tert-butyl ethyl ether, and toluene) were analyzed in groundwater from 49 of the 50 wells. Water from seven sites had detections for fuel compounds; all concentrations were below MCL. The results provided assurance of water quality and a valuable baseline to evaluate future trends of fuel constituents as the region is further developed. Probability maps were developed from logistic regression models to examine the likelihood that nitrate concentrations in groundwater exceeded specified levels. Susceptibility analysis examined relations between mid-level (5.0 mg/L) nitrate concentrations and climatic, hydrologic, and geologic variables; the significant variables were identified as depth to groundwater, soil organic matter, and soil water storage to 25-centimeter (cm) depth. The vulnerability assessments included natural factors driving susceptibility but also human factors related to land use and septic use. Vulnerability to low-level (2.5 mg/L) nitrate was related to depth to groundwater, septic zoning, and soil organic matter. The results highlighted that septic zoning affected low-level nitrate concentrations. Vulnerability to mid-level (5.0 mg/L) nitrate was examined using all 50 samples and also with two data outliers removed, which showed relatively high nitrate concentrations but also anomalous water chemistry or were located beyond the primary study area. Vulnerability to mid-level (5.0 mg/L) nitrate using all 50 samples was related to depth to groundwater, land use, septic use within a 500-meter (m) radius, soil water storage to a 25-cm depth, soil organic matter, and whether a location was within the agricultural region. The mid-level (5.0 mg/L) vulnerability model using 48 samples (two outliers removed) produced the best overall fit and was related to the same variables as when using all samples except septic use. The results for mid-level vulnerability provided additional support that septic use was associated with low levels of nitrate in the groundwater. Soil properties and land use were identified as the main drivers of moderate nitrate concentrations. Probabilities of exceeding low-level nitrate concentrations were high in most areas with the lowest probabilities usually to the northwest along thin geologic deposits in the upper part of the basin. The results of this investigation offer the foundational information needed for developing best management practices to mitigate nitrate contamination, basic concepts on water quality to aid public education, and information to guide regulatory measures if policy makers determine this is warranted. Science-based decision making will require continued monitoring and analysis of water quality in the future.

Colorado↗

Small creatures, big challenge: A review of current knowledge and assessment of research needs to inform water withdrawal management for the Duck River's freshwater mussel fauna

The Duck River is a remarkable national resource, boasting outstanding biodiversity (Ahlstedt et al. 2017). It contains 151 fish species, over 60 mussel species, and 22 snail species, many of which are federally listed (Etnier and Starnes 1993; Johnson et al. 2013; Ahlstedt et al. 2017; Womble and Rosenberger 2025a). The Duck River watershed is at a pivotal time in its management and ecological history due to significant growth in human population and economic activity. The consequent increase in surface water demand presents challenges for sustainable water resource management. The Tennessee Department of Environment and Conservation (TDEC) has permitted or proposed to permit eight water utilities to withdraw over 77 million gallons per day (mgd) from the Duck River, representing > 40% increase from previously permitted amounts (USFWS 2025). On November 20, 2024, Tennessee Governor Bill Lee signed Executive Order No 108 to ensure long-term stability of the Duck River watershed through balancing economic growth, water resource management, and environmental conservation. Some primary components of this plan are the creation of a Comprehensive Regional Drought Management Plan, the development of a Habitat Conservation Plan, and the establishment of the Duck River Watershed Planning Partnership (DRWPP). Although the planning partnership is not focused on a sole aspect of the Duck River, but the watershed as a whole, the mussel assemblages of the Duck River have been a particular focus due to their importance in providing ecosystem services, sensitivity to water quality, conservation status, and sensitivity to periodic drought conditions in this system. With multiple state and federal agencies involved in managing the Duck River’s unique water resources, a focused review on how future water management could affect the freshwater mussel communities in the Duck River will contribute to this process. Our objectives of this review are to: 1. Review the effects of changing flow regimes on mussels, with emphasis on the effect of droughts/low flows, highlighting key processes/factors affecting mussels. 2. Highlight critical data gaps that could allow future, evidence-based water management decisions in this system. 3. Propose research topics to address critical data gaps and enable evidence-based water management decisions that consider the needs of freshwater mussels. Freshwater mussels are a critical component of riverine ecosystems, providing ecosystem services including water filtration, nutrient cycling, and habitat provision (Vaughn et al. 2004; Haag 2012) that benefit human communities. Environmental flows, also referred to as ecological or e-flows, refer to the quantity, timing, and quality of water flows necessary to maintain the ecological health of rivers, wetlands, and other aquatic ecosystems (Acreman and Dunbar 2004; Acreman 2016). For freshwater mussels, environmental flows include maintaining a flow regime and associated water quality regimes that are favorable for both juveniles and adults while also seasonally supporting host fish and the survival of settling glochidia (i.e., larval mussels) in suitable environments (Gates et al. 2015). Freshwater mussels are adapted to the natural flow regime of their native streams, including seasonal variability in flows. As sedentary filter feeders, mussels require adequate water flow to provide oxygen and food (Allen and Vaughn 2009); however, they also require periods of low flow to increase access to fish hosts and to ensure juveniles are not displaced (Gates et al 2015). Mussels are sensitive to water quality conditions, including pollutants, sedimentation, and artificial alterations of temperature and pH (Strayer and Malcom 2012). Mussel community characteristics, including presence, abundance, and species richness, generally increase with stream size (e.g., stream order); however, responses to changes in flow conditions are often species-specific, and the magnitude and timing of extreme flow conditions have more predictive power for mussel community response than general changes in flow (Lopez and Vaughn 2021). Complex hydraulic variables, such as relative shear stress and Reynolds number, can be strong predictors of mussel community responses because they combine simple hydraulic variables (e.g., velocity, depth) with stream channel characteristics, such as substrate composition and dynamics. Droughts directly and indirectly affect mussels by influencing their physiology, behavior, reproduction, and ecosystem roles. Mass mussel die-offs during droughts and low water conditions are widely documented (Vaughn et al. 2015; Sousa et al. 2018; Dubose et al. 2019). Droughts can also lead to a shift in mussel community composition, with drought-tolerant species becoming more dominant while more sensitive species experience declines (Galbraith et al, 2010; Lopez et al. 2022; Vaughn and Atkinson 2025). Droughts affect mussels through direct mortality from stranding as well as physiological stress from altered water temperature and quality, reduced habitat and food availability and quality, increased predation and competition, disruption of reproductive cycles, and increased susceptibility to disease and pathogens. Although the effects of droughts, water withdrawals, and low flow conditions on mussels are an increasing area of study, there is a general paucity of data-driven consensus regarding optimal mitigation approaches and strategies (Cushway et al. 2024). Long-term management decisions regarding water withdrawals, particularly during extended or multi-year drought conditions, may consider factors beyond direct mortality such as the sublethal effects of drought and low flow conditions on freshwater mussels for a comprehensive understanding of the effects of prolonged low flows on freshwater mussels and aquatic communities.

Tennessee↗

Population and habitat analyses for greater sage-grouse (Centrocercus urophasianus) in the bi-state distinct population segment—2018 update

Executive Summary The Bi-State Distinct Population Segment (Bi-State DPS) of greater sage-grouse ( Centrocercus urophasianus , hereinafter “sage-grouse”) represents a genetically distinct and geographically isolated population that straddles the border between Nevada and California. The primary threat to these sage-grouse populations is the expansion of single-leaf pinyon ( Pinus monophylla ) and Utah juniper ( Juniperus osteosperma ) into sagebrush ecosystems, which fragments and reduces population connectivity and survival. Other important threats include low water availability during brood-rearing, particularly during drought, and increased predation by common ravens ( Corvus corax ), a generalist predator often associated with anthropogenic resource subsidies. Although the Bi-State DPS occurs at high elevations relative to sage-grouse range-wide, changes in historical wildfire cycles and the conversion of native shrubs to invasive annual grasslands still threaten these populations. The Bi-State DPS has undergone multiple federal status assessments and associated litigation. For example, in October of 2013, the Bi-State DPS was proposed for listing as threatened under the Endangered Species Act of 1973 by the U.S. Fish and Wildlife Service (USFWS), then withdrawn in April 2015. The withdrawal decision was challenged, and in May 2018, a Federal district court ordered the withdrawal decision to be vacated, and USFWS was required to re-open the October 2013 listing evaluation. In response, the U.S. Geological Survey (USGS), with State and Federal collaborators, embarked on a multipronged analysis to provide current and best available science regarding population status of sage-grouse within the Bi-State DPS. Using data from a long-term monitoring program, we carried out four analytical study objectives, and here, we provide preliminary results of these analyses. First, we used integrated population modeling (IPM) to predict annual population abundance and annual finite rate of population change for the Bi-State DPS, as a whole, and for each subpopulation between 1995 and 2018. Because sage-grouse exhibit population cycles (periodic increases and decreases in abundance across approximately 6- to 10-year wavelengths), we estimated trends across three nested temporal scales that represent one (11 years), two (18 years), and three (24 years) complete population cycles. These estimates of relatively long-term averaged population change account for temporal (that is, interannual) variation. Our model predicted population abundance for the Bi-State DPS during 2018 at 3,305 individuals (2,247–4,683), with the majority occupying Bodie Hills and Long Valley. The model also predicted cyclic dynamics in abundance through time with evidence of 24-year population growth and slight trends of decline over the past 18 years. Specifically, across the Bi-State DPS as a whole, we estimated annual average at 0.99, 0.99, and 1.02 over the one, two, and three population cycles, which equated to a 10.5 percent, 16.6 percent decrease, and 60.0 percent increase in abundance over the 11-, 18-, and 24-year cycles. Estimated abundance in 2018 had not reached numbers lower than those predicted during 1995. However, we found spatial variation in population trends across the three cycles. Bodie Hills subpopulation comprised the greatest (1,521) and exhibited average annual greater than 1.0 across all periods resulting in average annual increases of 7 percent. This relatively large subpopulation has grown 5 times larger than what was predicted in 1995 while experiencing cyclical dynamics within that period. Conversely, other smaller subpopulations within the Bi-State DPS exhibited average annual equal to or less than 1.0 resulting in estimated 10-year risks of extirpation ranging from 2.0 to 76.1 percent. In general, evidence of decline among smaller subpopulations was greatest for the most recent period (2008–18) compared to a period that encompassed three full population cycles (24-year). This difference coincides with an intense period of drought that began in 2012. For comparative purposes as part of this first objective, we conducted a similar analysis for populations of sage-grouse within Nevada and California but outside the Bi-State DPS. We developed a region-wide and distance-weighted IPM using lek count from Nevada Department of Wildlife (NDOW) and California Department of Fish and Wildlife (CDFW) databases and with telemetry data collected by USGS across 12 sage-grouse subpopulations. Our models predicted similar patterns in population cycling outside the Bi-State DPS but with much stronger evidence of long-term declines across 24 years. Specifically, median averaged across each year of the 11-, 18-, and 24-year periods resulted in average annual values of 0.94, 0.97, and 0.99, respectively. These values equate to 41.0 percent, 38.5 percent, and 21.3 percent declines over the corresponding periods. Second, we used lek count data in a state-space modeling framework to compare trends in population abundance across different spatial scales (that is, leks versus Bi-State DPS). This hierarchical framework allowed us to disentangle declines associated with climate conditions as opposed to other local level factors that might signal the need for management intervention. Specifically, we identified 7 leks that were both declining and recently decoupled from larger spatial scale trends, typically governed by climatic conditions (referred to as soft or hard signals). The goal of this analysis was to provide an early warning system that might have implications for conservation actions at local scales. Third, we developed phenological (spring, summer–fall, and winter) and reproductive life stage (nesting, early brood-rearing, and late-brood rearing) based resource selection functions using various environmental covariates. We report rankings of variable importance for each season and life stage, developed maps of habitat selection indices (HSI), binned categories representing low, moderate, and high classes of quality (where any category greater than or equal to low indicated selected habitat) for each phenological season and life stage, and produced composite maps by selected phenological and reproductive stage to estimate annual habitat. Fourth, we used for each lek within the Bi-State DPS to carry out a spatial analysis that quantified substantial changes in the distribution of occupied habitat across long- (24-year) and short- (11-year) term periods. Owing to differences among available datasets, the long-term analysis primarily reflected spatial shifts among subpopulations comprising the majority of the Bi-State DPS (that is, Bodie Hills and Long Valley) while the short-term analysis also quantified changes among subpopulations along the periphery. Over long and short-term periods, the overall distribution of occupied habitat (as measured by 99 percent utilization distributions intersecting any quantified habitat) was reduced by 20,573 ha and 55,492 ha, respectively. Occupied core areas (as measured by 50 percent utilization distributions intersecting any quantified habitat) over long-term periods were solely located in Bodie Hills and Long Valley. Although nearly all subpopulations experienced contractions in occupied overall and core distribution, Bodie Hills experienced spatial expansion that occurred with concomitant spatial contraction at Long Valley over both periods. Subpopulations at the northern (Pine Nuts), central (Sagehen) and southern (White-Mountains) extents of the Bi-State DPS also experienced spatial contraction over the short-term period. These findings, coupled with those of population trends, indicate long-term patterns in redistribution of sage-grouse from Long Valley and peripheral subpopulations to Bodie Hills. That is, sage-grouse subpopulations at the periphery are declining while the largest population at the core is increasing, which could have meaningful impacts on overall metapopulation persistence. We provide evidence for loss of occupied habitat (reduced distribution) given local extirpation of subpopulations. Fifth, we calculated percentages of selected phenological, life stage, and annual habitat that each subpopulation contributed to the Bi-State DPS. We then intersected these maps with a composite estimate of occupied habitat from the fourth objective and calculated percentages of selected habitat likely occupied by sage-grouse that each subpopulation contributed to the Bi-State DPS. These values provide evidence for loss of occupied habitat and subsequent reductions in spatial distribution given reductions in abundance and, in some cases, extirpation of leks within subpopulations. Lastly, we carried out an initial in-depth analysis of selection for irrigated pastures and wet meadows during the brood-rearing stage for the Long Valley subpopulation. We chose this subpopulation because it represents a population core, representing 26.5 percent of total sage-grouse within the Bi-State DPS, and has exhibited long-term declines in abundance and distribution. This subpopulation is highly sensitive to precipitation and other factors that influence water availability. Models predicted higher use of the interior portions of irrigated pastures and wet meadows during late brood-rearing period, which represented a potentially risky use of habitat that was exacerbated during periods of low moisture (for example, drought, reduced water delivery, or both). Sage-grouse typically used edges of riparian areas and pastures, largely because the interior of these mesic areas consisted of considerably less overhead concealment cover (for example, shrubs) that likely resulted in a higher risk of mortality. We found that a lack of water delivery to pastures in the form of overwinter precipitation or diversion ditches increased the movements of sage-grouse to the interior of pastures. Although further investigation of water delivery impacts on chick survival are warrented, our initial findings regarding resource selection may explain recent declines in population growth at Long Valley.

California, Nevada↗

FLUXNET-CH4: A global, multi-ecosystem database and analysis of methane seasonality from freshwater wetlands

Methane (CH 4 ) emissions from natural landscapes constitute roughly half of global CH 4 contributions to the atmosphere, yet large uncertainties remain in the absolute magnitude and the seasonality of emission quantities and drivers. Eddy covariance (EC) measurements of CH 4 flux are ideal for constraining ecosystem-scale CH 4 emissions due to quasi-continuous and high-temporal-resolution CH 4 flux measurements, coincident carbon dioxide, water, and energy flux measurements, lack of ecosystem disturbance, and increased availability of datasets over the last decade. Here, we (1) describe the newly published dataset, FLUXNET-CH 4 Version 1.0, the first open-source global dataset of CH 4 EC measurements (available at https://fluxnet.org/data/fluxnet-ch4-community-product/ , last access: 7 April 2021). FLUXNET-CH 4 includes half-hourly and daily gap-filled and non-gap-filled aggregated CH 4 fluxes and meteorological data from 79 sites globally: 42 freshwater wetlands, 6 brackish and saline wetlands, 7 formerly drained ecosystems, 7 rice paddy sites, 2 lakes, and 15 uplands. Then, we (2) evaluate FLUXNET-CH 4 representativeness for freshwater wetland coverage globally because the majority of sites in FLUXNET-CH 4 Version 1.0 are freshwater wetlands which are a substantial source of total atmospheric CH 4 emissions; and (3) we provide the first global estimates of the seasonal variability and seasonality predictors of freshwater wetland CH 4 fluxes. Our representativeness analysis suggests that the freshwater wetland sites in the dataset cover global wetland bioclimatic attributes (encompassing energy, moisture, and vegetation-related parameters) in arctic, boreal, and temperate regions but only sparsely cover humid tropical regions. Seasonality metrics of wetland CH 4 emissions vary considerably across latitudinal bands. In freshwater wetlands (except those between 20 ∘ S to 20 ∘ N) the spring onset of elevated CH 4 emissions starts 3 d earlier, and the CH 4 emission season lasts 4 d longer, for each degree Celsius increase in mean annual air temperature. On average, the spring onset of increasing CH 4 emissions lags behind soil warming by 1 month, with very few sites experiencing increased CH 4 emissions prior to the onset of soil warming. In contrast, roughly half of these sites experience the spring onset of rising CH 4 emissions prior to the spring increase in gross primary productivity (GPP). The timing of peak summer CH 4 emissions does not correlate with the timing for either peak summer temperature or peak GPP. Our results provide seasonality parameters for CH 4 modeling and highlight seasonality metrics that cannot be predicted by temperature or GPP (i.e., seasonality of CH 4 peak). FLUXNET-CH 4 is a powerful new resource for diagnosing and understanding the role of terrestrial ecosystems and climate drivers in the global CH 4 cycle, and future additions of sites in tropical ecosystems and site years of data collection will provide added value to this database. All seasonality parameters are available at https://doi.org/10.5281/zenodo.4672601 (Delwiche et al., 2021). Additionally, raw FLUXNET-CH 4 data used to extract seasonality parameters can be downloaded from https://fluxnet.org/data/fluxnet-ch4-community-product/ (last access: 7 April 2021), and a complete list of the 79 individual site data DOIs is provided in Table 2 of this paper.

Earth System Science Data↗

Nitrous oxide emission from denitrification in stream and river networks

Nitrous oxide (N 2 O) is a potent greenhouse gas that contributes to climate change and stratospheric ozone destruction. Anthropogenic nitrogen (N) loading to river networks is a potentially important source of N 2 O via microbial denitrification that converts N to N 2 O and dinitrogen (N 2 ). The fraction of denitrified N that escapes as N 2 O rather than N 2 (i.e., the N 2 O yield) is an important determinant of how much N 2 O is produced by river networks, but little is known about the N 2 O yield in flowing waters. Here, we present the results of whole-stream 15 N-tracer additions conducted in 72 headwater streams draining multiple land-use types across the United States. We found that stream denitrification produces N 2 O at rates that increase with stream water nitrate (NO 3 − ) concentrations, but that <1% of denitrified N is converted to N 2 O. Unlike some previous studies, we found no relationship between the N 2 O yield and stream water NO 3 − . We suggest that increased stream NO 3 − loading stimulates denitrification and concomitant N 2 O production, but does not increase the N 2 O yield. In our study, most streams were sources of N 2 O to the atmosphere and the highest emission rates were observed in streams draining urban basins. Using a global river network model, we estimate that microbial N transformations (e.g., denitrification and nitrification) convert at least 0.68 Tg·y −1 of anthropogenic N inputs to N 2 O in river networks, equivalent to 10% of the global anthropogenic N 2 O emission rate. This estimate of stream and river N 2 O emissions is three times greater than estimated by the Intergovernmental Panel on Climate Change. Humans have more than doubled the availability of fixed nitrogen (N) in the biosphere, particularly through the production of N fertilizers and the cultivation of N-fixing crops ( 1 ). Increasing N availability is producing unintended environmental consequences including enhanced emissions of nitrous oxide (N 2 O), a potent greenhouse gas ( 2 ) and an important cause of stratospheric ozone destruction ( 3 ). The Intergovernmental Panel on Climate Change (IPCC) estimates that the microbial conversion of agriculturally derived N to N 2 O in soils and aquatic ecosystems is the largest source of anthropogenic N 2 O to the atmosphere ( 2 ). The production of N 2 O in agricultural soils has been the focus of intense investigation (i.e., >1,000 published studies) and is a relatively well constrained component of the N 2 O budget ( 4 ). However, emissions of anthropogenic N 2 O from streams, rivers, and estuaries have received much less attention and remain a major source of uncertainty in the global anthropogenic N 2 O budget. Microbial denitrification is a large source of N 2 O emissions in terrestrial and aquatic ecosystems. Most microbial denitrification is a form of anaerobic respiration in which nitrate (NO 3 − , the dominant form of inorganic N) is converted to dinitrogen (N 2 ) and N 2 O gases ( 5 ). The proportion of denitrified NO 3 − that is converted to N 2 O rather than N 2 (hereafter referred to as the N 2 O yield and expressed as the mole ratio) partially controls how much N 2 O is produced via denitrification ( 6 ), but few studies provide information on the N 2 O yield in streams and rivers because of the difficulty of measuring N 2 and N 2 O production in these systems. Here we report rates of N 2 and N 2 O production via denitrification measured using whole-stream 15 NO 3 − -tracer experiments in 72 headwater streams draining different land-use types across the United States. This project, known as the second Lotic Intersite Nitrogen eXperiment (LINX II), provides unique whole-system measurements of the N 2 O yield in streams. Although N 2 O emission rates have been reported for streams and rivers ( 7 , 8 ), the N 2 O yield has been studied mostly in lentic freshwater and marine ecosystems, where it generally ranges between 0.1 and 1.0%, although yields as high as 6% have been observed ( 9 ). These N 2 O yields are low compared with observations in soils (0–100%) ( 10 ), which may be a result of the relatively lower oxygen (O 2 ) availability in the sediments of lakes and estuaries. However, dissolved O 2 in headwater streams is commonly near atmospheric equilibrium and benthic algal biofilms can produce O 2 at the sediment–water interface, resulting in strong redox gradients more akin to those in partially wetted soils. Thus, streams may have variable and often high N 2 O yields, similar to those in soils ( 11 ). The N 2 O yield in headwater streams is of particular interest because much of the NO 3 − input to rivers is derived from groundwater upwelling into headwater streams. Furthermore, headwater streams compose the majority of stream length within a drainage network and have high ratios of bioreactive benthic surface area to water volume ( 12 ).

Proceedings of the National Academy of Sciences of↗