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MetaIPM: Placing integral projection models into a metapopulation framework

Metapopulation models include spatial population dynamics such as dispersion and migration between subpopulations. Integral projection models (IPMs) can include demographic rates as a function of size. Traditionally, metapopulation models do not included detailed populaiton models such as IPMs. In some situations, both local population dynamics (e.g. size-based survival) and spatial dynamics are important. We present a Python package, MetaIPM , which places IPMs into a metapopulation framework, and allow users to readily construct and apply these models that combine local population dynamics within a metapopulation framework. MetaIPM includes an IPM for each subpopulation that is connected to other subpopulations via a metapopulation movement model. These movements can include dispersion, migration or other patterns. The IPM can include for size-specific demographic rates (e.g. survival, recruitment) as well as management actions, such as length-based harvest (e.g. gear specific capture sizes, varying slot limits across political boundaries). The model also allows for changes in metapopulation connectivity between locations, such as a fish passage ladders to enhance movement or deterrents to reduce movement. Thus, resource managers can use MetaIPM to compare different management actions such as the harvest gear type (which can be length-specific) and harvest locations. We demonstrate how MetaIPM may be applied to inform managers seeking to limit the spread of an invasive species in a system with important metapopulation dynamics. Specifically, we compared removal lengths (all length fish versus longer fish only) for an invasive fish population in a fragmented, inland river system. MetaIPM allowed users to compare the importance of harvesting source populations away from the invasion front, as well as species at the invasion front. The model would also allow for future comparisons of different deterrent placement locations in the system. Moving beyond our example system, we describe how MetaIPM can be applied to other species, systems and management approaches. The MetaIPM packages includes Jupyter Notebooks documenting the package as well as a second set of JupyterNotebooks showing the application of the package to our example system.

Methods in Ecology and Evolution↗

Carbon stocks across a chronosequence of thinned and unmanaged red pine ( Pinus resinosa ) stands

Forests function as a major global C sink, and forest management strategies that maximize C stocks offer one possible means of mitigating the impacts of increasing anthropogenic CO 2 emissions. We studied the effects of thinning, a common management technique in many forest types, on age-related trends in C stocks using a chronosequence of thinned and unmanaged red pine ( Pinus resinosa ) stands ranging from 9 to 306 years old. Live tree C stocks increased with age to a maximum near the middle of the chronosequence in unmanaged stands, and increased across the entire chronosequence in thinned stands. C in live understory vegetation and C in the mineral soil each declined rapidly with age in young stands but changed relatively little in middle-aged to older stands regardless of management. Forest floor C stocks increased with age in unmanaged stands, but forest floor C decreased with age after the onset of thinning around age 40 in thinned stands. Deadwood C was highly variable, but decreased with age in thinned stands. Total ecosystem C increased with stand age until approaching an asymptote around age 150. The increase in total ecosystem C was paralleled by an age-related increase in total aboveground C, but relatively little change in total belowground C. Thinning had surprisingly little impact on total ecosystem C stocks, but it did modestly alter age-related trends in total ecosystem C allocation between aboveground and belowground pools. In addition to characterizing the subtle differences in C dynamics between thinned and unmanaged stands, these results suggest that C accrual in red pine stands continues well beyond the 60–100 year management rotations typical for this system. Management plans that incorporate longer rotations and thinning in some stands could play an important role in maximizing C stocks in red pine forests while meeting other objectives including timber extraction, biodiversity conservation, restoration, and fuel reduction goals.

Ecological Applications↗

Ecosystem vs. community recovery 25 years after grass invasions and fire in a subtropical woodland

Despite a large body of research documenting invasive plant impacts, few studies have followed individual invaded sites over decades to observe how they change, and none have contrasted how compositional impacts from invasion compare to ecosystem-process impacts over a multi-decadal time-scale. Using direct measurements of plant density and composition and of ecosystems processes, we evaluate how ecosystem structure, above-ground net primary production (ANPP), and above-ground and soil nutrient pools compare over 25 years since fire and C4 grass invasions disrupted seasonally dry Hawaiian woodlands. We compare structure and function between primary woodland that has never burned and is largely native species-dominated, with sites that had been the same woodland type but burned in alien-grass-fuelled fires in the 1970s and 1980s. The sites have not experienced fires since 1987. We report here that woody plant composition and structure continue to be dramatically changed by the initial invasions and fires that occurred 25 years ago and invaders continue to dominate in burned sites. This is reflected in continued low plant carbon pools in burned compared to unburned sites. Yet ANPP and N storage, which were dramatically lower in the initial decade after invasive-grass fuelled fires, have increased and are now indistinguishable from values measured in intact woodlands. Soil carbon pools were resilient to both invasion and fire initially and over time. Above-ground net primary production has recovered because of invasion of burned sites by a non-native N-fixing tree rather than because of recovery of native species. This invasive N-fixing tree is unlikely to return C storage of the invaded sites to those of unburned woodland because of its tissue and growth characteristics and its interactions with invasive grasses. It does not facilitate native species but rather promotes a persistent invasive grass/N-fixer savanna. Synthesis. We conclude that fire, an unusual disturbance in this system, has perpetuated the dominance of these sites by invasive species and that despite the dramatic recovery of above-ground net primary production and N pools, the ecosystem continues to be in a distinctly different state than the pre-fire, pre-Melinis community. Thus, despite the absence of further disturbance (fire), there is no evidence that succession towards the original ecosystem is occurring. The fact that N pools and above-ground net primary production recover because of a new invader (Morella faya), highlights the unpredictability of ecosystem trajectories in the face of altered regional species pools.

Hawaii↗

Modeling soil porewater salinity in mangrove forests (Everglades, Florida, USA) impacted by hydrological restoration and a warming climate

Hydrology is a critical driver controlling mangrove wetlands structural and functional attributes at different spatial and temporal scales. Yet, human activities have negatively affected hydrology, causing mangrove diebacks and coverage loss worldwide. In fact, the assessment of mangrove water budgets, impacted by natural and human disturbances, is limited due to a lack of long-term data and information that hinders our understanding of how changes in hydroperiod and salinity control mangrove productivity and spatial distribution. In this study, we implemented a mass balance-based hydrological model (RHYMAN) that explicitly considers groundwater discharge in the Shark River estuary (SRE, southwestern Everglades) located in a karstic geomorphic setting and influenced by regional hydrological restoration. We used long-term hydroperiod and porewater salinity (PWS) datasets obtained from 2004 to 2016 for model calibration and validation and to determine spatiotemporal variability in water levels and PWS at three riverine mangrove sites (downstream, SRS-6; midstream, SRS-5; upstream, SRS-4) along SRE. Model results agree with a distinct PWS pattern along the estuarine salinity gradient where the highest PWS occurs at SRS-6 (mean: 25, range: 22–30 ppt), followed by SRS-5 (17, 14–25 ppt) and SRS-4 (5, 3–13 ppt). A commensurate increase in PWS over a thirteen-year period indicates a long-term reduction in freshwater inflow coupled with sea-level rise (SLR). Increasing freshwater scenario simulation results show a significant reduction (17–27%) in PWS along the estuary in contrast with a high SLR scenario when salinity increases up to 1.1 to 2.5 times that of control values. Model results show that freshwater inflow and SLR are key drivers controlling mangrove wetlands PWS in this karstic coastal region. Given its relatively simple structure, this mass balance-based hydrological model could be used in other environmental settings to evaluate potential habitat and regime shifts due to changes in hydrology and PWS under regional hydrological restoration management.

Florida↗

A regionally varying habitat model to inform management for greater sage-grouse persistence across their range

Identifying habitat needs for species with large distributions is challenging because species-habitat associations may vary across scales and regions (spatial nonstationarity). Furthermore, management efforts often cross jurisdictional boundaries, complicating the development of cohesive conservation strategies among management entities. The greater sage-grouse ( Centrocercus urophasianus ) is a rapidly declining species that spans 11 U.S. states and responds to habitat conditions across a wide range of spatial scales and regions. Allowing for regional variance in species-habitat associations and suitability predictions could systematically identify important habitats at levels relevant to management. We collaboratively developed a model with Bureau of Land Management (BLM) biologists that: (1) evaluated the scale of effect for different environmental covariates; (2) accounted for regional differences in population-level responses; and (3) predicted probabilities of persistence across the U.S. occupied range. We modeled range-wide lek persistence data (6615 communal breeding sites classified as active or inactive) as a function of environmental covariates. Environmental covariates included sagebrush cover, pinyon-juniper cover, topography, precipitation, point and line disturbance densities, and landscape configuration metrics. Our model treated habitat assessment areas – regionally delineated by BLM biologists – as random intercepts and slopes that allowed for geographic variation in species-habitat associations and predicted probabilities of lek persistence. Our final model indicated support for 12 environmental covariates predicting lek persistence at scales extending between 1- to 15-km radii from lek centers, and a covariate measuring distance to the occupied range boundary. Five of these covariates showed significant regionally varying responses: sagebrush clumpiness (a measure of habitat aggregation), pinyon-juniper cover, point disturbance of anthropogenic features such as energy infrastructure and communication towers, elevation, and a topographic index associated with mesic habitats. This spatial nonstationarity indicates unitary range-wide recommendations, or rules-of-thumb with respect to their effects on lek persistence, may be problematic for these environmental conditions. For covariates that did not include random slopes, and which were potentially amenable to management actions, we found that leks were predicted to become extirpated when sagebrush cover fell below 9.6 % (summarized at the 3.2-km radius extent), and the proportion of classified sagebrush habitat fell below 0.7 (1-km). We produced a continuous predictive probability surface of lek persistence which we binned based on model sensitivity thresholds to produce habitat quality categories. The highest quality habitat (capturing 50 % of active leks) covered 25.5 % of the occupied range, while the combined lowest through highest quality habitats (capturing 95 % of active leks) covered 65.0 %. Accommodating regional environmental differences in models that are relevant to habitat management planning will help ensure their applicability to targeted goals. Continuous collaboration between modelers and land managers early in the modeling process increases the likelihood of this outcome.

Global Ecology and Conservation↗

The noise is the signal: Spatio-temporal variability of production and productivity in high elevation meadows in the Sierra Nevada mountain range of North America

There are expectations that increasing temperatures will lead to significant changes in structure and function of montane meadows, including greater water stress on vegetation and lowered vegetation production and productivity. We evaluated spatio-temporal dynamics in production and productivity in meadows within the Sierra Nevada mountain range of North America by: (1) compiling Landsat satellite data for the Normalized Difference Vegetation Index (NDVI) across a 37-year period (1985–2021) for 8,095 meadows >2,500 m elevation; then, (2) used state-space models, changepoint analysis, geographically-weighted regression (GWR), and distance-decay analysis (DDA) to: (a) identify meadows with decreasing, increasing or no trends for NDVI; (b) detect meadows with abrupt changes (changepoints) in NDVI; and (c) evaluate variation along gradients of latitude, longitude, and elevation for eight indices of temporal dynamics in annual production (mean growing season NDVI; MGS) and productivity (rate of spring greenup; RSP). Meadows with no long-term change or evidence of increasing NDVI were 2.6x more frequent as those with decreasing NDVI (72% vs. 28%). Abrupt changes in NDVI were detected in 48% of the meadows; they occurred in every year of the study and with no indication that their frequency had changed over time. The intermixing of meadows with different temporal dynamics was a consistent pattern for monthly NDVI and, especially, the eight annual indices of MGS and RSP. The DDA showed temporal dynamics in pairs of meadow within a few 100 m of each other were often as different as those hundreds of kilometers apart. Our findings point strongly toward a great diversity of temporal dynamics in meadow production and productivity in the SNV. The heterogeneity in spatial patterns indicated that production and productivity of meadow vegetation is being driven by interplay among climatic, physiographic and biotic factors at basin and meadow scales. Thus, when evaluating spatio-temporal dynamics in condition for many high elevation meadow systems, what might often be considered “noise” may provide greater insight than a “signal” embedded within a large amount of variability.

California↗

Changes in vegetation in northern Alaska under scenarios of climate change, 2003-2100: Implications for climate feedbacks

Assessing potential future changes in arctic and boreal plant species productivity, ecosystem composition, and canopy complexity is essential for understanding environmental responses under expected altered climate forcing. We examined potential changes in the dominant plant functional types (PFTs) of the sedge tundra, shrub tundra, and boreal forest ecosystems in ecotonal northern Alaska, USA, for the years 2003–2100. We compared energy feedbacks associated with increases in biomass to energy feedbacks associated with changes in the duration of the snow-free season. We based our simulations on nine input climate scenarios from the Intergovernmental Panel on Climate Change (IPCC) and a new version of the Terrestrial Ecosystem Model (TEM) that incorporates biogeochemistry, vegetation dynamics for multiple PFTs (e.g., trees, shrubs, grasses, sedges, mosses), multiple vegetation pools, and soil thermal regimes. We found mean increases in net primary productivity (NPP) in all PFTs. Most notably, birch ( Betula spp.) in the shrub tundra showed increases that were at least three times larger than any other PFT. Increases in NPP were positively related to increases in growing-season length in the sedge tundra, but PFTs in boreal forest and shrub tundra showed a significant response to changes in light availability as well as growing-season length. Significant NPP responses to changes in vegetation uptake of nitrogen by PFT indicated that some PFTs were better competitors for nitrogen than other PFTs. While NPP increased, heterotrophic respiration ( R H ) also increased, resulting in decreases or no change in net ecosystem carbon uptake. Greater aboveground biomass from increased NPP produced a decrease in summer albedo, greater regional heat absorption (0.34 ± 0.23 W·m −2 ·10 yr −1 [mean ± SD]), and a positive feedback to climate warming. However, the decrease in albedo due to a shorter snow season (−5.1 ± 1.6 d/10 yr) resulted in much greater regional heat absorption (3.3 ± 1.24 W·m −2 ·10 yr −1 ) than that associated with increases in vegetation. Through quantifying feedbacks associated with changes in vegetation and those associated with changes in the snow season length, we can reach a more integrated understanding of the manner in which climate change may impact interactions between high-latitude ecosystems and the climate system.

Alaska↗

Plant-herbivore-hydroperiod interactions: effects of native mammals on floodplain tree recruitment

Floodplain plant–herbivore–hydroperiod interactions have received little attention despite their potential as determinants of floodplain structure and functioning. We used five types of exclosures to differentially exclude small-, medium-, and large-sized mammals from accessing Fremont cottonwood ( Populus deltoides Marshall subsp. wizlizenii (Watson) Eckenwalder) seedlings and saplings growing naturally on four landform types at an alluvial reach on each of two rivers, the Green and Yampa, in Colorado and Utah. The two study reaches differed primarily as a result of flow regulation on the Green River, which began in 1962. Landforms were a rarely flooded portion of the alluvial plain, geomorphically active slow- and fast-water channel margin sites on the Yampa reach, and an aggrading side channel on the Green. Small-mammal live-trapping and observational data indicated that, with minor exceptions, the kinds of mammals eating cottonwood within each reach were identical. We monitored condition and fates of individual cottonwood plants from October 1993 through the 1997 growing season. Differences in survival and growth were noted both within and between reaches, and both due to, and independent of, mammalian herbivory. Comparisons of cottonwood growth and survivorship among exclosures and between exclosures and controls indicated that a small mammal, Microtus montanus, reduced seedling and sapling survivorship at the Green River reach, but to a lesser extent (seedlings) or not at all (saplings) on the Yampa reach. In contrast, reductions in sapling height increment attributable to medium- and large-sized herbivores were detected only at the Yampa site. We suggest that these differences are a result of (1) flow regulation allowing Microtus populations to escape the mortality normally accompanying the large, snowmelt-driven spring flood, as well as regulation promoting a herbaceous understory favorable to voles, and (2) greater browsing pressure from overwintering deer and elk at the Yampa reach, unrelated to flow regulation. Within areas used by foraging beaver, the probability of a sapling being cut by beaver was similar on the two reaches. This study suggests that changes in riparian plant–herbivore relationships due to shifts in river hydrology may be a common and important consequence of river regulation.

Ecological Applications↗

Homogenization of soil seed bank communities by fire and invasive species in the Mojave Desert

Soil seed banks help maintain species diversity through temporal storage effects and function as germination pools that can optimize fitness across varying environmental conditions. These characteristics promote the persistence of native plant communities, yet disturbances such as fire and associated invasions by non-native species can disrupt these reserves, fundamentally altering successional trajectories. This may be particularly true in deserts, where native plant communities are less adapted to fire. While studies of fire effects on desert plant communities are not uncommon, information regarding the short- and long-term effects of fire on seed banks is less available. To better understand the influence of fire and invasive species on desert seed banks, we investigated soil seed bank biodiversity from 30 wildfires that burned between 1972 and 2010 across the Mojave Desert ecoregion of North America. We assessed how characteristics of fire regimes (frequency, time since fire, and burn severity) interacted with climate and invasive plants on measures of α-, β-, and γ-diversities. Because β-diversity is a direct measure of community variability and reveals important information about biodiversity loss, we further examined the nestedness and turnover components of β-diversity. Mean α- and γ-diversities were generally higher for burned locations than in unburned reference sites, however individual fire variables had little influence on patterns of seed bank diversity. Burned area seed banks tended to be dominated by non-native invasive species, primarily two grasses, ( Bromus rubens , Bromus tectorum ), as well as an invasive forb ( Erodium cicutarium ). The most striking pattern we observed was a collective sharp decline in α-, β-, and γ-diversities with increased invasive species dominance, indicating the homogenization of seed bank communities with the colonization of invasive species after fire. Evidence of homogenization was further supported by reduced turnover and increased nestedness in burn areas compared to reference areas indicating potential biodiversity loss. Our findings highlight how biological processes such as plant invasions can combine with disturbance from fire to alter patterns of seed bank composition and diversity in desert ecosystems.

Frontiers in Ecology and Evolution↗

Simulation of groundwater budgets and travel times for watersheds on the north shore of Long Island Sound, with implications for nitrogen-transport studies

Aquatic systems in and around the Long Island Sound (LIS) provide a variety of ecological and economic benefits, but in some areas of the LIS, aquatic ecosystems have become degraded by excess nitrogen. A substantial fraction of the nitrogen inputs to the LIS are transported through the groundwater-flow system. Because groundwater travel times in surficial aquifers can exceed 100 years, multiyear lags are introduced between inputs at the water table in recharge areas and discharge to inland or coastal receiving waters. The U.S. Geological Survey, in cooperation with the Connecticut Department of Energy and Environmental Protection and the U.S. Environmental Protection Agency’s Long Island Sound Study, developed a steady-state groundwater model of the watersheds draining from the northern shore of the LIS for the purpose of calculating groundwater budgets and travel times to coastal waters. The model was developed by using the MODFLOW–NWT software and existing spatial data on aquifers, river networks, land-surface altitudes, land cover, groundwater recharge, and water use. Coastal waters were delineated on the basis of the National Wetland Inventory; all non-coastal waters were collectively termed “inland waters.” A coarse-resolution model was calibrated by using the PEST++ software, long-term records of water levels in 65 wells, stream altitudes from 477 streams, base-flow records for 14 streamgages that are relatively unaffected by withdrawals, and error metrics based on incorrectly simulated flooding and incorrectly simulated dry streams. The calibrated values were used in a fine-resolution model in which the mean absolute residuals were 4.5 meters for groundwater levels, 1.3 meters for stream altitudes, and 7,200 cubic meters per day (2.9 cubic feet per second) for base flow. About 89 percent of the terrestrial cells were correctly simulated with the water table below land surface, and nearly 90 percent of the cells representing streams were correctly simulated as having the water table above the stream bottom. Together, these metrics suggest that this model is robust for simulating regional-scale groundwater patterns. Simulated groundwater budgets were compiled for the entire study area, for each HUC12 (Hydrologic Unit Code no. 12) watershed and its adjacent coastal waters, if applicable, within the study area, and for 14 coastal-embayment watersheds. Most groundwater (90.6 percent of inflows) discharged to inland waters, with smaller fractions to coastal waters (7.0 percent) and well withdrawals (2.4 percent). When computed for HUC12 watersheds with coastal discharge, the portions of groundwater discharging to coastal waters ranged from 0.02 to 66 percent of groundwater outflows, with a median of 13 percent. Within priority-embayment watersheds, the portions of groundwater discharging to coastal waters ranged from 2 to 56 percent, with a median of 15 percent. Groundwater travel times also were simulated for the entire study area, for each HUC12 watershed and its adjacent coastal waters, if applicable, within the study area and for 14 priority coastal embayments. Within the entire study area, the median groundwater travel time was 1.9 years, with an interquartile range of 0.1 to 5.9 years. Sensitivity analysis of groundwater travel times within a subbasin in the study area indicates that the travel times are a function of the grid resolution, with coarser grids resulting in shorter median travel times. Travel times for groundwater discharging to coastal waters were similar to travel times for groundwater discharging to inland waters, with a median of 1.9 years. Median travel times for the HUC12 watersheds ranged from 0.9 to 53.5 years, with a median of 1.8 years. Among HUC12 watersheds that include coastal areas, travel times for groundwater discharging to coastal waters ranged from less than 1 to 61.6 years, with a median of 2.8 years. The HUC12 watersheds with the longest simulated travel times were in the urban area near New York City where the model performance is less accurate. Median travel times for groundwater discharging to coastal waters within the priority-embayment watersheds ranged from less than 1 to 18.6 years, with a median of 2.3 years. A more focused analysis was conducted for the Niantic River watershed to demonstrate the applicability of the regional model to local-scale nitrogen-transport analyses by using nitrogen-input and -attenuation rates from literature sources. Nitrogen inputs were estimated by using land-cover-based loading factors, and attenuation was estimated by using attenuation factors based on geologic zones and soil properties. Based on this analysis, groundwater transports an estimated 22,000 kilograms of nitrogen per year (2.9 kilograms of nitrogen per hectare per year) to streams, rivers, and coastal waters within the Niantic River watershed. Approximately 36 percent of discharging nitrogen is from atmospheric-deposition sources, 38 percent is from fertilizers, and 26 percent is from septic systems. Most of the groundwater-transported nitrogen (88 percent) discharges first to streams and rivers, with only 12 percent discharging directly to coastal waters. Travel times for groundwater-transported nitrogen ranged from less than 1 day to more than 100 years, with a median of 1.6 years.

Connecticut, New York, Rhode Island↗

Predicting 21st-century polar bear habitat distribution from global climate models

Projections of polar bear (Ursus maritimus) sea ice habitat distribution in the polar basin during the 21st century were developed to understand the consequences of anticipated sea ice reductions on polar bear populations. We used location data from satellitecollared polar bears and environmental data (e.g., bathymetry, distance to coastlines, and sea ice) collected from 1985 to 1995 to build resource selection functions (RSFs). RSFs described habitats that polar bears preferred in summer, autumn, winter, and spring. When applied to independent data from 1996 to 2006, the RSFs consistently identified habitats most frequently used by polar bears. We applied the RSFs to monthly maps of 21st-century sea ice concentration projected by 10 general circulation models (GCMs) used in the Intergovernmental Panel of Climate Change Fourth Assessment Report, under the A1B greenhouse gas forcing scenario. Despite variation in their projections, all GCMs indicated habitat losses in the polar basin during the 21st century. Losses in the highest-valued RSF habitat (optimal habitat) were greatest in the southern seas of the polar basin, especially the Chukchi and Barents seas, and least along the Arctic Ocean shores of Banks Island to northern Greenland. Mean loss of optimal polar bear habitat was greatest during summer; from an observed 1.0 million km 2 in 1985-1995 (baseline) to a projected multi-model mean of 0.32 million km 2 in 2090-2099 (-68% change). Projected winter losses of polar bear habitat were less: from 1.7 million km 2 in 1985-1995 to 1.4 million km 2 in 2090-2099 (-17% change). Habitat losses based on GCM multi-model means may be conservative; simulated rates of habitat loss during 1985-2006 from many GCMs were less than the actual observed rates of loss. Although a reduction in the total amount of optimal habitat will likely reduce polar bear populations, exact relationships between habitat losses and population demographics remain unknown. Density and energetic effects may become important as polar bears make long-distance annual migrations from traditional winter ranges to remnant high-latitude summer sea ice. These impacts will likely affect specific sex and age groups differently and may ultimately preclude bears from seasonally returning to their traditional ranges.

Ecological Monographs↗

Evaluating management alternatives for Wyoming elk feedgrounds in consideration of chronic wasting disease

Executive Summary The authors used decision and modeling analyses to evaluate management alternatives for a decision on whether to permit Cervus canadensis (elk) feeding on two sites on Bridger-Teton National Forest, Dell Creek and Forest Park. Supplemental feeding of elk could increase the transmission of chronic wasting disease (CWD) locally and disease spread regionally, potentially impacting elk populations over time with wider implications for Odocoileus hemionus (mule deer) and Odocoileus virginianus (white-tailed deer) populations and hunting, tourism, and regional revenue. Supplemental feeding is thought to improve overwinter elk survival and reduce the commingling of elk with cattle during months when brucellosis transmission risk is highest. We worked with the U.S. Department of Agriculture Forest Service to identify their fundamental objectives and associated performance metrics related to this feedground decision. We then developed disease and habitat selection models to quantify the effect of four management alternatives on select performance metrics. The four alternatives were to continue to permit feeding, phaseout permits to feed in three years, permit feeding on an emergency basis, or stop permitting feeding. In this report, we present methods and summarized results on disease and habitat selection models and summaries of other performance metrics analyzed by BIO-WEST, Inc. and Cirrus Ecological Solutions as part of an Environmental Impact Statement. Data from Wyoming Game and Fish Department (WGFD) supported the assumption that supplemental elk feeding allows for larger elk populations in a region. We documented that herd units (HU) without feedgrounds had 23 percent lower densities of elk per area of winter range when compared against HUs with feedgrounds, after accounting for differences in sightability of elk during counts on and off feedgrounds. Thus, throughout our analyses, we assumed feedground closures would reduce elk carrying capacity resulting in an average decline of previously fed elk population segments by 23 percent (5th and 95th percentiles = [11 percent, 35 percent]) by year 20. Most of that decline occurred within the first few years after a feedground ceases to operate. We used a panel of CWD experts to help estimate CWD trans-mission in fed and unfed elk population segments. In aggregate, the expert panel estimated that median values of direct and indirect transmission of CWD are expected to be 1.9 and 4 times higher, respectively, in fed elk populations compared to unfed elk. We used these disease transmission estimates in combination with local elk demographic rates and carrying capacity estimates to project disease and population dynamics. In year 20, we predicted CWD prevalence would increase to 42 percent (5th and 95th percentiles = [29 percent, 55 percent]), and 13 percent (5th and 95th percentiles = [4 percent, 26 percent]) on average for fed and unfed elk population segments, respectively, given a starting prevalence of 1.6 percent. The prevalence estimates for the unfed elk population segments are in the range of previous observations of CWD in elk in the western United States. The average CWD prevalence from 2016 to 2018 in the unfed elk population of Wind Cave National Park in South Dakota was 18 percent overall but up to 30 percent in some regions (Sargeant and others, 2021). Meanwhile, CWD prevalence in the Iron Mountain and Laramie Peak elk herds in Wyoming from 2016 to 2018 was 14 percent and 7 percent, respectively, despite being present since at least 2002 (Wyoming Game and Fish Department, 2020b). From 2016 to 2020, elk that were fed at Dell Creek and Forest Park constituted on average 12–20 percent of the total elk on their respective HUs. As a result, the differences between management alternatives are modest when considering the closure of only one feedground on a HU. The no feeding alternative for Forest Park resulted in a CWD prevalence of 17 percent (SD = 7 percent) in the Afton HU compared to 20 percent (SD = 7 percent) with continued feeding by year 20. In the Upper Green River HU, no feeding on Dell Creek resulted in a CWD prevalence of 27 percent (SD = 6 percent) compared to 30 percent (SD = 5 percent) with continued feeding. In terms of disease-associated mortality, we predicted the closure of Forest Park and Dell Creek feedgrounds would reduce the total number of CWD mortalities by 9 percent in the Upper Green River HU and 26 percent in the Afton HU during the 20-year timespan. Our spatial analyses predicted that management alternative effects vary by HU as a function of private property and other wildlife winter ranges proximity relative to feedground location. The predicted number of elk abortions on private land, as a proxy for brucellosis risk to cattle, may increase by 8–21 percent in the absence of feeding at Dell Creek and Forest Park. Eight feedgrounds are located on Bridger-Teton National Forest, all of which have permits that have expired or will expire prior to 2028. In addition, WGFD could change their management of feedgrounds given new information; therefore, we also assessed the cumulative effects of continued feeding, phaseout, and no feeding management alternatives across five HUs south of Jackson, Wyoming (Afton HU, Fall Creek HU, Piney HU, Pinedale HU, and Upper Green River HU). These five HUs ranged from about 41 to 85 percent of the elk herd using feedgrounds, which corresponded to a CWD prevalence at year 20 of 23–34 percent if all feedgrounds in those five HUs remained open relative to 12 to 14 percent if all feedgrounds were closed. We predicted feedground closures may result in immediate reductions in population size relative to alternatives that continue feeding (for example, continued feeding and emergency feeding alternatives); however, over longer periods of time, CWD-associated mortality leads to larger population reductions. The no feeding alternative resulted in higher elk population sizes compared to the continued feeding alternative after about 10 years of implementation. Delayed action under a phaseout alternative resulted in increasing the CWD prevalence to 20 percent relative to 12 to 14 percent, on average, without feeding on HUs with a large population of fed elk such as the Upper Green River HU. Summarizing our cumulative results across all five of the analyzed HUs, we predicted continued feeding will lead to fewer elk by year 20 (mean = 8,300, standard deviation [SD] = 740) compared to no feeding at U.S. Department of Agri-culture Forest Service sites (10,700, SD = 890). The closure of all feedgrounds was projected to result in the largest elk populations at year 20 (12,500, SD = 980). No feeding at all sites also resulted in the largest cumulative harvest of 57,700 (SD = 2,600) compared to 51,100 (SD = 3,800) for continued feeding at all current feedground sites on the five HUs. Continued feeding also resulted in the lowest brucellosis costs to producers ($194,600, SD = $11,500) compared to no feeding on all feedgrounds ($243,000, SD = $13,700). Assuming moderate reductions in hunter interest because of increasing CWD prevalence in elk, we predicted that no feeding resulted in regional revenues generated by hunting activities of $190 million (SD = $10 million) compared to $173 million (SD = $10 million) for continued feeding over the 20-year timeframe. Recent CWD detections in mule deer and elk in Grand Teton National Park has elevated the importance of the cur-rent decision on whether, and how, to permit elk feeding on Dell Creek and Forest Park and the management of the other feedgrounds. Aggressive male harvest has slowed, but not stopped, the increasing prevalence of CWD in mule deer (Conner and others, 2021). It is unclear whether harvest management can be an effective tool to slow the spread of CWD in elk. There are also no effective treatments or vaccines for CWD, and it is unlikely that any will be developed that can be easily deployed in the near future. Thus, reducing artificial aggregations is one of the few management approaches suggested by the Western Association of Fish and Wildlife Agencies (Almberg and others, 2017). Future surveillance and monitoring can be designed to resolve uncertainties that can improve future decision-making. If feedgrounds close, research could quantify elk population reductions in the absence of feeding, the redistribution of fed elk to other places, or the consequences of elk movement on private property. If feedgrounds remain open, research could assess how rapidly CWD spreads in artificial aggregations of elk; however, surveillance programs would need to be designed with sufficient power to detect initial changes of CWD prevalence. Delaying action on feedground management was projected to be costly. Results of the phaseout alternative relative to the no feeding alternative suggested a 3-year delay was enough for substantial long-term changes in CWD prevalence. The long-term persistence of infectious CWD prions in the environment suggests that feedground management decisions may have long-lasting consequences. Our results indicated tradeoffs in the ability of a management agency to achieve all their objectives, and all management alternatives resulted in significant reductions in elk population size. This report contains the foundational elements for formal decision analysis methods, which can be implemented to help decision makers transparently evaluate the consequences of decision alternatives and identify the set of actions that best achieve agency and stakeholder priorities.

Wyoming↗

Looking ahead, guided by the past: The role of U.S. national parks in amphibian research and conservation

Protected areas like national parks are essential elements of conservation because they limit human influence on the landscape, which protects biodiversity and ecosystem function. The role of national parks in conservation, however, often goes far beyond limiting human influence. The U.S. National Park Service and its system of land units contribute substantively to conservation by providing protected lands where researchers can document trends in species distributions and abundances, examine characteristics important for generating these trends, and identify and implement conservation strategies to preserve biodiversity. We reviewed the contribution of U.S. national parks to amphibian research and conservation and highlight important challenges and findings in several key areas. First, U.S. national parks were instrumental in providing strong support that amphibian declines were real and unlikely to be simply a consequence of habitat loss. Second, research in U.S. national parks provided evidence against certain hypothesized causes of decline, like UV-B radiation, and evidence for others, such as introduced species and disease. However, describing declines and identifying causes contributes to conservation only if it leads to management; importantly, U.S. national parks have implemented many conservation strategies and evaluated their effectiveness in recovering robust amphibian populations. Among these, removal of invasive species, especially fishes; conservation translocations; and habitat creation and enhancement stand out as examples of successful conservation strategies with broad applicability. Successful management for amphibians is additionally complicated by competing mandates and stakeholder interests; for example, past emphasis on increasing visitor enjoyment by introducing fish to formerly fishless lakes had devastating consequences for many amphibians. Other potential conflicts with amphibian conservation include increasing development, increased risk of introductions of disease and exotic species with increased visitation, and road mortality. Decision science and leveraging partnerships have proven to be key components of effective conservation under conflicting mandates in national parks. As resource managers grapple with large-scale drivers that are outside local control, public-private partnerships and adaptive strategies are increasing in importance. U.S. national parks have played an important role in many aspects of identifying and ameliorating the amphibian decline crisis and will continue to be essential for the conservation of amphibians in the future.

Ecological Indicators↗

Food web conceptual model

This chapter describes a general model of food webs within tidal wetlands and represents how physical features of the wetland affect the structure and function of the food web. This conceptual model focuses on how the food web provides support for (or may reduce support for) threatened fish species. This model is part of a suite of conceptual models designed to guide monitoring of restoration sites throughout the San Francisco Estuary (SFE), but particularly within the Sacramento-San Joaquin Delta (Delta) and Suisun Marsh. The conceptual models have been developed based on the Delta Regional Ecosystem Restoration Implementation Plan (DRERIP) models, and are designed to aid in the identification and evaluation of monitoring metrics for tidal wetland restoration projects. Many tidal restoration sites in the Delta are being constructed to comply with environmental regulatory requirements associated with the operation of the Central Valley Project and State Water Project. These include the Biological Opinions for Delta Smelt (Hypomesus transpacificus) and salmonids (U.S. Fish and Wildlife Service 2008; National Marine Fisheries Service 2009), and the Incidental Take Permit for Longfin Smelt (Spirinchus thaleichthyes) (California Department of Fish and Wildlife 2009). These regulatory requirements are based on the hypothesis that the decline of listed fish species is due in part to a decline in productivity of the food web (phytoplankton and zooplankton in particular) or alterations in the food web such that production is consumed by other species in the Estuary (Sommer et al. 2007; Baxter et al. 2010; Brown et al. 2016a). Intertidal wetlands and shallow subtidal habitat can be highly productive, so restoring areas of tidal wetlands may result in a net increase in productivity that will provide food web support for these fish species. However, other factors such as invasive bivalves that reduce phytoplankton and zooplankton biomass and invasive predatory fishes that may compete with or prey upon listed fishes can limit the utility of tidal wetlands for food web support (Lucas and Thompson 2012; Herbold et al. 2014). This model utilizes information from the previous DRERIP models for Delta food webs (Durand 2008) and tidal wetlands (Kneib et al. 2008), an updated DRERIP model (Durand 2015), and the State of BayDelta Science 2016 review of recent Delta food web literature (Brown et al. 2016a).

Interagency Ecological Program Technical Report↗

Vegetation, fuels, and fire-behavior responses to linear fuel-break treatments in and around burned sagebrush steppe: Are we breaking the grass-fire cycle?

Background Linear fuel breaks are being implemented to moderate fire behavior and improve wildfire containment in semiarid landscapes such as the sagebrush steppe of North America, where extensive losses in perennial vegetation and ecosystem functioning are resulting from invasion by exotic annual grasses (EAGs) that foster large and recurrent wildfires. However, fuel-break construction can also pose EAG invasion risks, which must be weighed against the intended fire-moderation benefits of the treatments. We investigated how shrub reductions (mowing, cutting), pre-emergent EAG-herbicides, and/or drill seedings of fire-resistant perennial bunchgrasses (PBGs) recently applied to create a large fuel-break system affected native and exotic plant abundances and their associated fuel loading and predicted fire behavior. Results In heavily EAG-invaded areas, herbicides reduced EAG and total herbaceous cover without affecting PBGs for 2–3 years and reduced predicted fire behavior for 1 year (from the Fuel Characteristic Classification System). However, surviving post-herbicide EAG cover was still > 30%, which was sufficient fuel to exceed the conventional 1.2-m-flame length (FL) threshold for attempting wildfire suppression with hand tools. In less invaded shrubland, shrub reduction treatments largely reduced shrub cover and height by ~ half without increasing EAGs, but then redistributed the wood to ground level and increased total herbaceous cover. Herbicides and/or drill seeding after shrub reductions did not affect EAG cover, although drill seedings increased PBG cover and exotic forbs (e.g., Russian thistle). Fire behavior was predicted to be moderated in only one of the many yearly observations of the various shrub-reduction treatment combinations. Over all treatments and years, FLs were predicted to exceed 1.2 m in 13% of simulations under average (11 km h −1 ) or high (47 km h −1 ) wind speed conditions and exceed the 3.4-m threshold for uncontrollable fire in 11% of simulations under high-wind speeds only. Conclusions Predicted fire-moderation benefits over the first 4 years of fuel break implementation were modest and variable, but, generally, increases in EAGs and their associated fire risks were not observed. Nonetheless, ancillary evidence from shrublands would suggest that treatment-induced shifts from shrub to herbaceous fuel dominance are expected to improve conditions for active fire suppression in ways not readily represented in available fire models.

Idaho, Oregon↗

Multiple pathways for woody plant establishment on floodplains at local to regional scales

1. The structure and functioning of riverine ecosystems is dependent upon regional setting and the interplay of hydrologic regime and geomorphologic processes. We used a retrospective analysis to study recruitment along broad, alluvial valley segments (parks) and canyon segments of the unregulated Yampa River and the regulated Green River in the upper Colorado River basin, USA. We precisely aged 811 individuals of Populus deltoides ssp. wislizenii (native) and Tamarix ramosissima (exotic) from 182 wooded patches and determined the elevation and character of the germination surface for each. We used logistic regression to relate recruitment events (presence or absence of cohort) to five flow and two weather parameters. 2. Woody plant establishment occurred via multiple pathways at patch, reach and segment scales. Recruitment occurred through establishment on (1) vertically accreting bars in the unregulated alluvial valley, (2) high alluvial floodplain surfaces during rare large flood events, (3) vertically accreting channel margin deposits in canyon pools and eddies, (4) vertically accreting intermittent/abandoned channels, (5) low elevation gravel bars and debris fans in canyons during multi-year droughts, and (6) bars and channels formed prior to flow regulation on the dammed river during controlled flood events. 3. The Yampa River's peak flow was rarely included in models estimating the likelihood that recruitment would occur in any year. Flow variability and the interannual pattern of flows, rather than individual large floods, control most establishment. 4. Regulation of the Green River flow since 1962 has had different effects on woody vegetation recruitment in canyons and valleys. The current regime mimics drought in a canyon setting, accelerating Tamarix invasion whereas in valleys the ongoing geomorphic adjustment of the channel, combined with reduced flow variability, has nearly eliminated Populus establishment . 5. A single year's flow or a particular pattern of flows over a sequence of years, whether natural or man-made, produces different recruitment opportunities in alluvial and canyon reaches, in diverse landforms within a particular river reach, and for Populus and Tamarix . The design of flows to restore riparian ecosystems must consider these multiple pathways and adjust the seasonal timing, magnitude and interannual frequency of flows to match the desired outcome.

Journal of Ecology↗

Accounting for non-random samples with distance sampling to estimate population density

A critical assumption of standard distance sampling is that sampling lines are located such that animals are uniformly distributed as a function of distance from the line. Failure to meet this assumption can introduce bias in the estimator. Many studies have used landscape features, such as roads or rivers, as lines, which can violate assumptions of distance sampling in two ways. First, animals may be attracted or repelled by the landscape feature due to human activity (e.g. along roads) or habitat characteristics associated with the feature (e.g. rivers). Second, sampling along landscape features may not be representative of the larger area of interest. We used auxiliary data to generalize the distance sampling estimator and relax assumptions of a uniform distribution of animals relative to distance from the line (i.e. density gradient) and to allow the distribution of animals to differ by habitat type. The generalized estimator provides unbiased estimates of density within the area sampled but may not be representative of the study area. To address the problem of landscape features providing unrepresentative sampling, we used a resource selection model to estimate the proportion of the population that occurred within the surveyed area to obtain an estimate of abundance for the desired area of inference. We demonstrate our modified distance sampling estimator using white-tailed deer ( Odocoileus virginianus ) in a 972-km 2 study area. We conducted infrared surveys of deer from roads to collect distance-to-transect data. We used locations of radio-collared deer to model the distribution of deer relative to the transects and to develop a resource selection model of deer based on distance to roads, habitat type, elevation and slope to account for roads being a non-representative sample of the study area. Synthesis and applications . When using landscape features as survey lines, the density gradient and deer distribution can introduce either positive or negative bias, which makes it impossible to assess the bias introduced without auxiliary data. The estimator we developed can improve precision because we obtained a better fit to distance observations and accounts for non-random placement of transects with minimal loss of precision.

Journal of Applied Ecology↗

Evidence that acidification-induced declines in plant diversity and productivity are mediated by changes in below-ground communities and soil properties in a semi-arid steppe

Anthropogenic acid deposition–induced soil acidification is one of the major threats to biodiversity, ecosystem functioning and services. Few studies, however, have explored in detail how above-ground changes in plant species richness and productivity resulting from soil acidification are mediated by effects on below-ground biota and soil properties. To increase our understanding of this linkage, we collected data on below- and above-ground communities and soil properties in a 3-year field experiment with seven levels of acid addition rate to build-up broad intensities of soil acidification in the semi-arid Inner Mongolian grassland. Acid addition directly elevated concentrations of soil Al 3+ ions, decreased the base cations Ca 2+ , Mg 2+ and Na + , and increased soil moisture and available phosphorus. Acid addition also appears to have altered the soil microbial community via changes in H + and Al 3+ ions and altered the nematode community via changes in H + ions and soil moisture. The observed changes in soil N availability (i.e. net N mineralization, NO 3 - -N and NH 4 + -N) could be explained by mediating changes in the H + and Al 3+ ions, microbial community (i.e. community structure, bacteria and fungi/bacteria as indicated by phospholipid fatty acids analysis) and the nematode community (i.e. total abundance, taxa richness and maturity index). Declines in plant species richness and productivity were greater at high intensities of soil acidification in the second sampling year than in the first sampling year. The changes in plant community observed were mostly explained by soil nutrient pathways (e.g. N availability or base mineral cations), which were in turn regulated by the soil microbial or nematode communities as well as by the direct effects of the increase in H + or Al 3+ ions. Synthesis. Our results suggest that the below-ground microbial and nematode communities are more sensitive to soil acidification than the plant communities are, and further that soil acidification–induced changes in plants are mediated by changes in below-ground communities and soil nutrients. These findings improve our understanding of the links between below- and above-ground communities in the Inner Mongolia grassland, especially in the context of anthropogenic acid enrichment.

Inner Mongolia↗

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