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Biological pathways of exposure and ecotoxicity values for uranium and associated radionuclides: Chapter D in Hydrological, geological, and biological site characterization of breccia pipe uranium deposits in Northern Arizona

This chapter compiles available chemical and radiation toxicity information for plants and animals from the scientific literature on naturally occurring uranium and associated radionuclides. Specifically, chemical and radiation hazards associated with radionuclides in the uranium decay series including uranium, thallium, thorium, bismuth, radium, radon, protactinium, polonium, actinium, and francium were the focus of the literature compilation. In addition, exposure pathways and a food web specific to the segregation areas were developed. Major biological exposure pathways considered were ingestion, inhalation, absorption, and bioaccumulation, and biota categories included microbes, invertebrates, plants, fishes, amphibians, reptiles, birds, and mammals. These data were developed for incorporation into a risk assessment to be conducted as part of an environmental impact statement for the Bureau of Land Management, which would identify representative plants and animals and their relative sensitivities to exposure of uranium and associated radionuclides. This chapter provides pertinent information to aid in the development of such an ecological risk assessment but does not estimate or derive guidance thresholds for radionuclides associated with uranium. Previous studies have not attempted to quantify the risks to biota caused directly by the chemical or radiation releases at uranium mining sites, although some information is available for uranium mill tailings and uranium mine closure activities. Research into the biological impacts of uranium exposure is strongly biased towards human health and exposure related to enriched or depleted uranium associated with the nuclear energy industry rather than naturally occurring uranium associated with uranium mining. Nevertheless, studies have reported that uranium and other radionuclides can affect the survival, growth, and reproduction of plants and animals. Exposure to chemical and radiation hazards is influenced by a plant’s or an animal’s life history and surrounding environment. Various species of plants, invertebrates, fishes, amphibians, reptiles, birds, and mammals found in the segregation areas that are considered species of concern by State and Federal agencies were included in the development of the site-specific food web. The utilization of subterranean habitats (burrows in uranium-rich areas, burrows in waste rock piles or reclaimed mining areas, mine tunnels) in the seasonally variable but consistently hot, arid environment is of particular concern in the segregation areas. Certain species of reptiles, amphibians, birds, and mammals in the segregation areas spend significant amounts of time in burrows where they can inhale or ingest uranium and other radionuclides through digging, eating, preening, and hibernating. Herbivores may also be exposed though the ingestion of radionuclides that have been aerially deposited on vegetation. Measured tissues concentrations of uranium and other radionuclides are not available for any species of concern in the segregation areas. The sensitivity of these animals to uranium exposure is unknown based on the existing scientific literature, and species-specific uranium presumptive effects levels were only available for two endangered fish species known to inhabit the segregation areas. Overall, the chemical toxicity data available for biological receptors of concern were limited, although chemical and radiation toxicity guidance values are available from several sources. However, caution should be used when directly applying these values to northern Arizona given the unique habitat and life history strategies of biological receptors in the segregation areas and the fact that some guidance values are based on models rather than empirical (laboratory or field) data. No chemical toxicity information based on empirical data is available for reptiles, birds, or wild mammals; therefore, the risks associated with uranium and other radionuclides are unknown for these biota.

Arizona↗

Quantifying ecological integrity of terrestrial systems to inform management of multiple-use public lands in the United States

The concept of ecological integrity has been applied widely to management of aquatic systems, but still is considered by many to be too vague and difficult to quantify to be useful for managing terrestrial systems, particularly across broad areas. Extensive public lands in the western United States are managed for diverse uses such as timber harvest, livestock grazing, energy development, and wildlife conservation, some of which may degrade ecological integrity. We propose a method for assessing ecological integrity on multiple-use lands that identifies the components of integrity and levels in the ecological hierarchy where the assessment will focus, and considers existing policies and management objectives. Both natural reference and societally desired environmental conditions are relevant comparison points. We applied the method to evaluate the ecological integrity of shrublands in Nevada, yielding an assessment based on six indicators of ecosystem structure, function, and composition, including resource- and stressor-based indicators measured at multiple scales. Results varied spatially and among indicators. Invasive plant cover and surface development were highest in shrublands in northwest and southeast Nevada. Departure from reference conditions of shrubland area, composition, patch size, and connectivity was highest in central and northern Nevada. Results may inform efforts to control invasive species and restore shrublands on federal lands in Nevada. We suggest that ecological integrity assessments for multiple-use lands be grounded in existing policies and monitoring programs, incorporate resource- and stressor-based metrics, rely on publicly available data collected at multiple spatial scales, and quantify both natural reference and societally desired resource conditions.

Nevada↗

A spatial model to prioritize sagebrush landscapes in the intermountain west (U.S.A.) for restoration

The ecological integrity of Sagebrush (Artemisia spp.) ecosystems in the Intermountain West (U.S.A.) has been diminished by synergistic relationships among human activities, spread of invasive plants, and altered disturbance regimes. An aggressive effort to restore Sagebrush habitats is necessary if we are to stabilize or improve current habitat trajectories and reverse declining population trends of dependent wildlife. Existing economic resources, technical impediments, and logistic difficulties limit our efforts to a fraction of the extensive area undergoing fragmentation, degradation, and loss. We prioritized landscapes for restoring Sagebrush habitats within the intermountain western region of the United States using geographic information system (GIS) modeling techniques to identify areas meeting a set of conditions based on (1) optimum abiotic and biotic conditions favorable for revegetation of Sagebrush; (2) potential to increase connectivity of Sagebrush habitats in the landscape to benefit wildlife; (3) location of population strongholds for Greater Sage-Grouse (Centrocercus urophasianus, a species of conservation concern); and (4) potential impediments to successful restoration created by Cheatgrass (Bromus tectorum, an invasive exotic annual grass). Approximately 5.8 million ha in southwestern Idaho, northern Nevada, and eastern Oregon met our criteria for restoring Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) and 5.1 million ha had high priority for restoring Mountain big sagebrush (A. tridentata ssp. vaseyana). Our results represent an integral component in a hierarchical framework after which site-specific locations for treatments can be focused within high-priority areas. Using this approach, long-term restoration strategies can be implemented that combine local-scale treatments and objectives with large-scale ecological processes and priorities. ?? 2008 Society for Ecological Restoration International.

Restoration Ecology↗

Drought resistance and resilience: The role of soil moisture–plant interactions and legacies in a dryland ecosystem

In many regions of the world, climate change is projected to reduce water availability through changes in the hydrological cycle, including more frequent and intense droughts, as well as seasonal shifts in precipitation. In water-limited ecosystems, such as drylands, lower soil water availability may exceed the adaptive capacity of many organisms, leading to cascading ecological effects during (concurrent effects) and after drought (legacy effects). The magnitude and duration of concurrent and legacy effects depends on drought intensity, duration and timing as well as the resistance and resilience of the ecosystem. Here, we investigated the effects of drought seasonality and plant community composition on two dominant perennial grasses, Achnatherum hymenoides (C 3 photosynthesis) and Pleuraphis jamesii (C 4 photosynthesis), in a dryland ecosystem. The experiment consisted of three precipitation treatments: control (ambient precipitation), cool-season drought (−66% ambient precipitation November–April) and warm-season drought (−66% ambient precipitation May–October), applied in two plant communities (perennial grasses with or without a large shrub, Ephedra viridis ) over a 3-year period. We examined the concurrent and legacy effects of seasonal drought on soil moisture, phenology and biomass. Drought treatments had strong concurrent and legacy effects on soil moisture, which impacted the phenology and biomass of the two grasses. Drought reduced growing season length by delaying green-up (cool-season drought) or advancing senescence (warm-season drought) and reduced biomass for both species. Biomass and phenology legacy effects from drought emerged in the second and third years of the experiment. While we observed differential sensitivity to drought legacies between the two grasses, we found limited evidence that shrub presence had interactive effects with the drought treatment. Synthesis . The results from this study highlight how abiotic and biotic legacies can develop and influence a community's resistance and resilience to subsequent droughts. When the frequency of repeated extreme events, such as recurring seasonal droughts, exceeds the capacity of organisms or ecosystems to recover (i.e. resilience), persistent drought legacies can reduce the resistance to subsequent drought events. Overall, these results highlight how drought legacies are a product of ecological resistance and resilience to past drought and can influence ecosystem vulnerability to future droughts.

Utah↗

Integrating observations and models to determine the effect of seasonally frozen ground on hydrologic partitioning in alpine hillslopes in the Colorado Rocky Mountains, USA

This study integrated spatially distributed field observations and soil thermal models to constrain the impact of frozen ground on snowmelt partitioning and streamflow generation in an alpine catchment within the Niwot Ridge Long-Term Ecological Research site, Colorado, USA. The study area was comprised of two contrasting hillslopes with notable differences in topography, snow depth and plant community composition. Time-lapse electrical resistivity surveys and soil thermal models enabled extension of discrete soil moisture and temperature measurements to incorporate landscape variability at scales and depths not possible with point measurements alone. Specifically, heterogenous snowpack thickness (~0–4 m) and soil volumetric water content between hillslopes (~0.1–0.45) strongly influenced the depths of seasonal frost, and the antecedent soil moisture available to form pore ice prior to freezing. Variable frost depths and antecedent soil moisture conditions were expected to create a patchwork of differing snowmelt infiltration rates and flowpaths. However, spikes in soil temperature and volumetric water content, as well as decreases in subsurface electrical resistivity revealed snowmelt infiltration across both hillslopes that coincided with initial decreases in snow water equivalent and early increases in streamflow. Soil temperature, soil moisture and electrical resistivity data from both wet and dry hillslopes showed that initial increases in streamflow occurred prior to deep soil water flux. Temporal lags between snowmelt infiltration and deeper percolation suggested that the lateral movement of water through the unsaturated zone was an important driver of early streamflow generation. These findings provide the type of process-based information needed to bridge gaps in scale and populate physically based cryohydrologic models to investigate subsurface hydrology and biogeochemical transport in soils that freeze seasonally.

Colorado↗

Linking modern pollen accumulation rates to biomass: Quantitative vegetation reconstruction in the western Klamath Mountains, NW California, USA

Quantitative reconstructions of vegetation abundance from sediment-derived pollen systems provide unique insights into past ecological conditions. Recently, the use of pollen accumulation rates (PAR, grains cm −2 year −1 ) has shown promise as a bioproxy for plant abundance. However, successfully reconstructing region-specific vegetation dynamics using PAR requires that accurate assessments of pollen deposition processes be quantitatively linked to spatially-explicit measures of plant abundance. Our study addressed these methodological challenges. Modern PAR and vegetation data were obtained from seven lakes in the western Klamath Mountains, California. To determine how to best calibrate our PAR-biomass model, we first calculated the spatial area of vegetation where vegetation composition and patterning is recorded by changes in the pollen signal using two metrics. These metrics were an assemblage-level relevant source area of pollen (aRSAP) derived from extended R-value analysis ( sensu Sugita, 1993) and a taxon-specific relevant source area of pollen (tRSAP) derived from PAR regression ( sensu Jackson, 1990). To the best of our knowledge, aRSAP and tRSAP have not been directly compared. We found that the tRSAP estimated a smaller area for some taxa (e.g. a circular area with a 225 m radius for Pinus ) than the aRSAP (a circular area with a 625 m radius). We fit linear models to relate PAR values from modern lake sediments with empirical, distance-weighted estimates of aboveground live biomass (AGL dw ) for both the aRSAP and tRSAP distances. In both cases, we found that the PARs of major tree taxa – Pseudotsuga, Pinus, Notholithocarpus , and TCT (Taxodiaceae, Cupressaceae, and Taxaceae families) – were statistically significant and reasonably precise estimators of contemporary AGL dw . However, predictions weighted by the distance defined by aRSAP tended to be more precise. The relative root-mean squared error for the aRSAP biomass estimates was 9% compared to 12% for tRSAP. Our results demonstrate that calibrated PAR-biomass relationships provide a robust method to infer changes in past plant biomass.

California, Oregon↗

South Florida wetlands ecosystem; biogeochemical processes in peat

The South Florida wetlands ecosystem is an environment of great size and ecological diversity (figs. 1 and 2). The landscape diversity and subtropical setting of this ecosystem provide a habitat for an abundance of plants and wildlife, some of which are unique to South Florida. South Florida wetlands are currently in crisis, however, due to the combined effects of agriculture, urbanization, and nearly 100 years of water management. Serious problems facing this ecosystem include (1) phosphorus contamination producing nutrient enrichment, which is causing changes in the native vegetation, (2) methylmercury contamination of fish and other wildlife, which poses a potential threat to human health, (3) changes in the natural flow of water in the region, resulting in more frequent drying of wetlands, loss of organic soils, and a reduction in freshwater flow to Florida Bay, (4) hypersalinity, massive algal blooms, and seagrass loss in parts of Florida Bay, and (5) a decrease in wildlife populations, especially those of wading birds. This U.S. Geological Survey (USGS) project focuses on the role of organic-rich sediments (peat) of South Florida wetlands in regulating the concentrations and impact of important chemical species in the environment. The cycling of carbon, nitrogen, phosphorus, and sulfur in peat is an important factor in the regulation of water quality in the South Florida wetlands ecosystem. These elements are central to many of the contamination issues facing South Florida wetlands, such as nutrient enrichment, mercury toxicity, and loss of peat. Many important chemical and biological reactions occur in peat and control the fate of chemical species in wetlands. Wetland scientists often refer to these reactions as biogeochemical processes, because they are chemical reactions usually mediated by microorganisms in a geological environment. An understanding of the biogeochemical processes in peat of South Florida wetlands will provide a basis for evaluating the effects on water quality of (1) constructing buffer wetlands to alleviate nutrient contamination and (2) replumbing the ecosystem to restore natural water flow. The results may also suggest new approaches for solving problems of contamination and water quality in these wetlands. A second focus of this project will be on the geochemical history of the South Florida ecosystem. Peat is a repository of the history of past environmental conditions in the wetland. Before effective action can be taken to correct many of the problems facing these wetlands, we must first study the biogeochemistry of the peat at depth in order to understand whether current problems are the result of recent human activity or are part of a long-term natural cycle. Coordination with other (USGS) projects for South Florida is ongoing. These projects are studying the biological history of the ecosystem by using pollen and shells buried in the peat, together with procedures for dating the peat at various depths, to develop an overall ecosystem history model, with emphasis on the last 100 years.

Fact Sheet↗

Remote sensing of land surface phenology

Remote sensing of land-surface phenology is an important method for studying the patterns of plant and animal growth cycles. Phenological events are sensitive to climate variation; therefore phenology data provide important baseline information documenting trends in ecology and detecting the impacts of climate change on multiple scales. The USGS Remote sensing of land surface phenology program produces annually, nine phenology indicator variables at 250 m and 1,000 m resolution for the contiguous U.S. The 12 year archive is available at http://phenology.cr.usgs.gov/index.php .

Fact Sheet↗

Conservation challenges emerging from free-roaming horse management: a vexing social-ecological mismatch

Horses have been associated with human societies for millennia, and for many have come to symbolize wildness, power, resilience, and freedom. Although equids were extirpated from North America 10 000-12 000 years ago, descendants of domestic horses now roam freely in the USA and 17 other countries across six continents. In landscape-scale and experimental investigations, free-roaming horses (Equus caballus) have been shown to induce numerous alterations to native-ecosystem components and processes through influences on soil, water, plants, native ungulates, small mammals, birds, reptiles, and other aspects of biodiversity. However, we argue that the management of free-roaming horses both in the U.S. and globally has been complicated by “socio-ecological mismatches.” These three mismatches stem from discrepancies between: 1) steady-state (horse-management) goals vs. a stochastically variable environment; 2) different methods to evaluate evidence, resolve conflicting beliefs, and achieve objectives, in scientific vs. political processes; and 3) differing spatial and temporal scales between policy and management, and values of science vs. values of the general public. Such mismatches arise from an inability to reconcile conflicting processes and functions in a social-ecological system, often reflecting differences in the spatio-temporal scales at which diverse components operate. Here, we describe three types of mismatches, and illustrate how the ecological dynamics of aridlands generally fit poorly with existing approaches to horse management and policy. Such mismatches complicate cost-effective management of free-roaming horses and the ecosystems they inhabit, and reduce the palette of potential solutions.

Conservation Letters↗

Using biodiversity metrics to guide conservation planning in altered tropical landscapes

Biodiversity metrics are frequently used to guide conservation planning because they can summarize biogeographical attributes of plant and animal communities quickly and at multiple scales. Attributes include habitat features of high conservation value, representativeness, and redundancy of biological communities. We conducted a rapid ecological assessment of resident avian species in the west-central mountainous region of Puerto Rico in 2015, a landscape dominated by coffee cultivation. We focused on this landscape because shade-grown and restored shade-grown coffee plantations offer an opportunity to complement protected habitat (e.g., reserves) to enhance species persistence. We used species richness, which tallies the number of unique species, and a quadratic entropy index of diversity, which incorporates interspecific taxonomic differentiation to evaluate species representativeness and redundancy across sun- and shade-grown coffee plantations and secondary forest. We surveyed 120 sites, calculating both metrics using species-specific occupancy probabilities estimated from community-level occupancy models. Species representativeness and redundancy were high as neither metric was able to discriminate among habitat types, possibly because plant communities were redundant, and the avian community was dominated by species adept at exploiting altered habitats. Similarly, we could not discriminate among avian communities modeling each biodiversity metric as a function of site-specific habitat covariates. Our findings and available knowledge on avian community demographics suggest that conservation strategies could couple protected habitat (e.g., reserves) and restored habitat (e.g., coffee plantations) to enhance species diversity and persistence across human-modified landscapes.

Caribbean Naturalist↗

Plant diversity shifts with drying and wetting cycles following removal of poplar plantations in Dongting Lake, China

Dongting Lake, the second largest lake in China, plays a crucial role in maintaining regional ecological security by providing key habitats for waterbirds. This study examined how hydrology and topography (ditches and mounds) together generate fine-scale wet–dry gradients that shape species establishment and successional trajectories. Over the past decade, vegetation restoration has been implemented through the removal of poplar plantations. We selected sites in West Dongting Lake representing different stages following poplar removal, including active plantations (Poplar), one-year (1Yr) and five-year (5Yr) restored sites, and natural floodplain reference sites. Analyses of seasonal water-level dynamics over several years revealed distinct hydrological regimes among restoration stages, with differences in inundation duration and amplitude corresponding to ditch–mound topography. Species diversity indicators showed that species richness was highest in Poplar (37 species) and lowest in Reference sites (14 species), with exotic species richness following the same trend. The lowest Cody index occurred between 1Yr and Reference sites, while the Jaccard similarity differed significantly among site pairs, with lower similarity observed in comparisons with the Reference site. Non-metric Multidimensional Scaling (NMDS) analysis indicated minimal overlap between Poplar and Reference sites, with 1Yr and 5Yr sites clustering together but remaining distinct from the Reference site. These results suggest that poplar cultivation and its associated mound-ditch topography have altered hydrological and community structure of the wetland, facilitating the persistence of exotic species. Natural recolonization is driven by the interaction between seasonal water-level dynamics and topography, which regulate inundation duration and wet–dry cycles. Effective restoration may require removal of poplar plantations and regrading of artificial ditches and mounds to restore natural hydrological connectivity to support vegetation recovery in these subtropical monsoonal wetlands.

Dongting Lake↗

Nutritional ecology of a fossorial herbivore: protein N and energy value of winter caches made by the northern pocket gopher, Thomomys talpoides

Northern pocket gophers ( Thomomys talpoides ) are fossorial herbivores that excavate belowground plant parts for food. In subalpine areas during autumn and winter, pocket gophers hoard plant parts in caches placed in or under snow. We examined the size and composition of 17 nival caches and tested the hypotheses that (i) cached food can provide complete energy and protein N sustenance during typical periods when burrowing is precluded by soil conditions, and (ii) cached food is a random sample of items encountered by burrowing gophers during tunnel excavation. Our data indicate that caches provide substantially more energy than protein in terms of a pocket gopher's daily maintenance requirements. Nevertheless, quantities stored are sufficient to allow individuals to endure commonly encountered adverse environmental conditions without entering negative energy or protein balance. Analysis of stomach contents and a comparison of cache composition to availability of plant species suggests that gophers consume high-protein items as they are encountered, and store low-protein items in caches.

Canadian Journal of Zoology↗

Development of a macrophyte-based index of biotic integrity for Minnesota lakes

Traditional approaches for managing aquatic resources have often failed to account for effects of anthropogenic disturbances on biota that are not directly reflected by chemical and physical proxies of environmental condition. The index of biotic integrity (IBI) is a potentially effective assessment method to integrate ecological, functional, and structural aspects of aquatic systems. A macrophyte-based IBI was developed for Minnesota lakes to assess the ability of aquatic plant communities to indicate environmental condition. The index was developed using quantitative point intercept vegetation surveys for 97 lakes that represent a range of limnological and watershed characteristics. We followed an approach similar to that used in Wisconsin to develop the aquatic macrophyte community index (AMCI). Regional adaptation of the AMCI required the identification of species representative of macrophyte communities in Minnesota. Metrics and scaling methods were also substantially modified to produce a more empirically robust index. Regression analyses indicated that IBI scores reflected statewide differences in lake trophic state (R2 = 0.57, F = 130.3, df = 1, 95, p < 0.005), agricultural (R2 = 0.51, F = 83.0, df = 1, 79, p < 0.005), urban (R2 = 0.22, F = 23.0, df = 1, 79, p < 0.005), and forested land uses (R2 = 0.51, F = 84.7, df = 1, 79, p < 0.005), and county population density (R2 = 0.14, F = 16.6, df = 1, 95, p < 0.005). Variance partitioning analyses using multiple regression models indicated a unique response of the IBI to human-induced stress separate from a response to natural lake characteristics. The IBI was minimally affected by differences in sample point density as indicated by Monte Carlo analyses of reduced sampling effort. Our analysis indicates that a macrophyte IBI calibrated for Minnesota lakes could be useful for identifying differences in environmental condition attributed to human-induced stress gradients. ?? 2010 Elsevier Ltd.

Ecological Indicators↗

The cost of self-defense: Browsing effects in the rare plant species Salix arizonica

Coevolution between plants and their animal predators has led to diverse defensive adaptations. Multiple theories of defense propose that there are resource allocation costs associated with producing chemical defenses. One leading hypothesis, optimal defense theory (ODT), suggests that natural selection will result in the allocation of resources to defenses that optimize the cost-to-benefit ratio between defense and other functional processes. The population decline of the rare subalpine wetland species, Arizona willow ( Salix arizonica ), has been attributed to various biotic and abiotic factors, with browsing from wild and domestic ungulates as a significant concern for at least three decades. In a field experiment using natural populations, we compare the relationship between phytochemical defense and height in Arizona willows with and without long-term protection from browsing via browse exclosures. Consistent with the predictions of ODT, individuals with physical protection from ungulate browsing for multiple years had significantly lower phenolic glycoside (PG) concentrations and increased plant height compared to unprotected individuals. A similar pattern was found across all individuals, whereby total PG concentration and height were negatively correlated. In a short-term experiment in natural populations, changes in levels of defense were not observed when plants received protection for only one growing season. The contrasting pattern of defense plasticity in response to long-term versus short-term physical protection suggests a differential plastic response in this long-lived species. Delayed reduction in PG concentration may serve as a benefit to avoid mismatches between environmental cues and responses. Our research sheds light on the intricate dynamics between plant-defense strategies, environmental pressures, and evolutionary adaptations in shaping plant–browser interactions.

Arizona, Colorado, New Mexico, Utah↗

Effects of water level, shade and time on germination and growth of freshwater marsh plants along a simulated successional gradient

1. We examined the effect of soil organic content (1.4, 3.6, 7.2% soil C), water level (+2, -1, -4 cm from soil surface) and duration (13 or 33 days) on 10 species that varied in abundance during succession in freshwater marshes. We also determined the effect of shade (0, 40, 80% shade) and soil organic content (1.4 and 7.2% soil C) on germination of six species over 62 days with water 0.5 cm below the soil surface. 2. Water level consistently affected species germination on both dates. Above-ground biomass was generally higher with increasing organic content of soil, but shade had little effect on germination or height. 3 The hydrologic zone in which species were found in the field was a good indicator of the response of germination to hydrology. Both early successional species and species wide-spread across the successional gradient show similar germination on all organic contents, while later successional species appear to germinate best at higher organic contents. 4. Successional changes in soils are capable of affecting plant community development, independent of disturbance.

Journal of Ecology↗

Effects of nutrient patches and root systems on the clonal plasticity of a rhizomatous grass

Clonal plant foraging has been examined primarily on individual clones exposed to resource-poor and resource-rich environments. We designed an experiment to examine the clonal foraging behavior of the rhizomatous grass Elymus lanceolatus ssp. lanceolatus under the influence of neighboring plant root systems in a heterogeneous nutrient environment. Individual Elymus clones were planted in large bins together with one of three neighboring grass species, Agropyron desertorum, Pseudoroegneria spicata, or Bromus tectorum, which differ in rooting density and growth activity. The position of Elymus clones was manipulated so rhizomes encountered a short-duration nutrient patch and subsequently root systems of the neighboring plants. Unexpectedly, the morphological plasticity of the perennial grass Elymus lanceolatus ssp. lanceolatus was influenced by the presence of the neighboring species much more than by the local nutrient enrichments, although nutrient patches did amplify some of the foraging responses. Elymus rhizomes branched readily and initiated large daughter plants as they encountered the low-density root systems of Pseudoroegneria. When Elymus encountered the fine, dense root systems of the annual Bromus, clonal expansion was initially reduced. Yet, after the short growing season of Bromus, Elymus resumed clonal expansion and produced several daughter plants. Elymus clones were most constrained by the fine, dense root systems of Agropyron desertorum. In this case, a few, long rhizomes avoided the densely rooted soil environment by growing aboveground as stolons crossing over the Agropyron tussocks. Elymus clonal biomass was largest in neighborhoods of Pseudoroegneria, intermediate in neighborhoods with Bromus, and smallest in neighborhoods with Agropyron. The latter were approximately half the size of those in the Pseudoroegneria environments. Elymus growth could not be explained by simple resource competition alone; other mechanisms must have been involved in the apparent differences in interference patterns of neighboring plants with Elymus.

Ecology↗

Faunal responses to fire in chaparral and sage scrub in California, USA

Impact of fire on California shrublands has been well studied but nearly all of this work has focused on plant communities. Impact on and recovery of the chaparral fauna has received only scattered attention; this paper synthesizes what is known in this regard for the diversity of animal taxa associated with California shrublands and outlines the primary differences between plant and animal responses to fire. We evaluated the primary faunal modes of resisting fire effects in three categories: 1) endogenous survival in a diapause or diapause-like stage, 2) sheltering in place within unburned refugia, or 3) fleeing and recolonizing. Utilizing these patterns in chaparral and sagescrub, as well as some studies on animals in other mediterranean-climate ecosystems, we derived generalizations about how plants and animals differ in their responses to fire impacts and their post fire recovery. One consequence of these differences is that variation in fire behavior has a much greater potential to affect animals than plants. For example, plants recover from fire endogenously from soil-stored seeds and resprouts, so fire size plays a limited role in determining recovery patterns. However, animals that depend on recolonization of burned sites from metapopulations may be greatly affected by fire size. Animal recolonization may also be greatly affected by regional land use patterns that affect colonization corridors, whereas such regional factors play a minimal role in plant community recovery. Fire characteristics such as rate of spread and fire intensity do not appear to play an important role in determining patterns of chaparral and sage scrub plant recovery after fire. However, these fire behavior characteristics may have a profound role in determining survivorship of some animal populations as slow-moving, smoldering combustion may limit survivorship of animals in burrows, whereas fast-moving, high intensity fires may affect survivorship of animals in above ground refugia or those attempting to flee. Thus, fire regime characteristics may have a much greater effect on postfire recovery of animal communities than plant communities in these shrubland ecosystems.

Fire Ecology↗

Ecological effects of pinyon-juniper removal in the Western United States—A synthesis of scientific research, January 2014–March 2021

Executive Summary Increasing density of pinyon ( Pinus spp. ) and juniper ( Juniperus spp. ) woodlands (hereinafter “pinyon-juniper”), as well as expansion of these woodlands into adjacent shrublands and grasslands, has altered ecosystem function and wildlife habitat across large areas of the interior western United States. Although there are many natural and human-caused drivers of woodland infilling and expansion, restoration of sagebrush ( Artemisia spp. ) habitat through removal of pinyon-juniper is considered an urgent management objective in many locations, particularly in support of sagebrush-dependent wildlife species of conservation concern. In December 2020, the Bureau of Land Management (BLM) established the Pinyon-Juniper Management Categorical Exclusion (PJCX) to expedite the regulatory process for pinyon-juniper removal projects on public lands, largely intended to benefit mule deer ( Odocoileus hemionus ) and greater sage-grouse ( Centrocercus urophasianus ) habitats. During final preparation of this report, the BLM discontinued use of the PJCX (as of November, 2022), but the pinyon-juniper tree removal techniques assessed in this report are commonly used and understanding their effects remains relevant to land use planning. To address areas of uncertainty relative to potential ecological effects of the PJCX, we conducted a review of the peer-reviewed science literature to better understand the likely responses of vegetation, environmental (for example, soils), and wildlife variables to specific tree removal techniques permitted by the PJCX. In brief, the PJCX permitted removal of trees by either manual cutting, mechanical cutting, or mastication; allowed certain methods to redistribute or remove resulting tree biomass after treatment; and prohibited broadcast burning, roadbuilding, removal of old-growth, and seeding of non-native species. Specifically, we conducted our review to address the following questions: How will PJCX removal techniques affect plant communities, soils, and abiotic resources? How do these pinyon-juniper removal techniques affect wildlife communities, including both woodland- and sagebrush-dependent species? What are the potential ecological implications of different pinyon-juniper removal treatment types and implementation strategies (for example, treatment sizes) over time? What are the most important gaps in our scientific understanding of how treatments might affect targeted ecosystems over space and time (for example, potential effects of climate change)? To answer these questions, we considered studies related to pinyon-juniper ecosystems, focusing on research that occurred over a large portion of the interior western United States that is the primary focus of the PJCX. We also focused on papers published from 2014 onward, to avoid excessive overlap with other recent reviews on pinyon-juniper management effects. Using strict criteria, including only considering research that tested responses for statistical significance, we identified 48 papers that primarily examined treatment effects on vegetation and other environmental variables (1,709 responses), and 11 papers that addressed effects on wildlife (132 responses). Responses to the PJCX-permitted treatments were summarized as either positive (that is, a significant increase), negative (that is, a significant decrease), or non-significant (that is, no significant difference). Responses were assigned to categories (for example, Native Annual Grass/Forb Abundance) and hierarchical treatment levels. We found that there were large proportions of non-significant responses among all categories combined, with roughly half or more of all responses non-significant (48 percent for wildlife, 60 percent for vegetation-environmental), comparable to other recent systematic reviews of pinyon-juniper treatment effects. However, we also found that when there were significant responses, some important trends potentially emerged. Important undesirable outcomes included far more positive than negative responses of exotic grass and forb abundance among nearly all treatment types. Cutting treatments were also more likely to decrease biocrust cover and microbial activity. Potentially beneficial outcomes included mostly positive responses among sagebrush obligate species, including more positive than negative responses for mule deer and sage-grouse. Some treatment types (for example, mastication) also resulted in more positive than negative responses for native grasses and forbs (although, non-significant responses were the majority). We also highlighted many limitations of this review, including how responses often come from few studies, and how some response-treatment category combinations lack adequate response data. Moreover, the existing research is often insufficient to address many key questions about treatment effects, largely owing to short time-scales and limited spatial extents of observations, which do not match the size of treatments being implemented by land managers, nor capture long-term, post-treatment ecological dynamics. We also identify a lack of research that addresses key interactions that could undermine restoration objectives, including potential effects of climate change and grazing on post-treatment environments. Thus, we emphasize the importance of integrating these factors into future pinyon-juniper treatment research, and we stress the need for use of monitoring programs and research studies that partake in data collection and analysis over long durations and broad spatial scales.

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