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Relative influence of flow regime, natural and anthropogenic environment on multidimensional stream fish diversity

The flow regime is considered a ‘master variable’ in riverine ecology because it directly influences stream geomorphology and biological communities. However, other environmental and anthropogenic factors have direct and synergistic effects with flow on fish diversity, complicating estimates of the flow regime's true importance. Moreover, most flow-ecology studies focus only on taxonomic diversity (i.e., species), without considering functional (trait-focused) or phylogenetic (evolution-focused) dimensions of diversity. In this study, we used linear regression with variation partitioning to parse out the independent and shared roles of the flow regime, physical environmental factors (e.g., soil characteristics) and the anthropogenic environment (e.g., developed land cover) for structuring multidimensional diversity of 365 stream fish communities in two biogeographic regions across South Carolina, USA. These variables explained between 8% and 18% of total variation of the local diversity. The flow regime contributed to diversity in all cases, but frequently covaried with physical and/or anthropogenic environmental variables. This covariation indicated that the independent role of flow would have been inflated if other environmental variables were not considered. The three dimensions of stream fish diversity were weakly correlated with one another and were associated with different environmental variables, indicating that each of them represents unique and complimentary facets of fish diversity. Accordingly, only considering the independent effects of instream flow on fish diversity may miss meaningful interactions between the flow regime and the other components of the environment that influence biodiversity patterns, leading to over simplified flow–ecology relationships.

South Carolina

The quality of our Nation’s waters--Ecological health in the Nation's streams, 1993-2005

This report summarizes a national assessment of the ecological health of streams done by the U.S. Geological Survey's (USGS) National Water-Quality Assessment Program (NAWQA). Healthy functioning stream ecosystems provide society with many benefits, including water purification, flood control, nutrient recycling, waste decomposition, fisheries, and aesthetics. The value to society of many of these benefits is substantial; for example, sportfishing in the United States generates an estimated annual economic output of $125 billion, including more than 1 million jobs (National Research Council, 2005; American Sportfishing Association, 2008). Continued monitoring and assessment of the Nation’s streams is needed to support informed decisions that will safeguard this important natural and economic resource. The quality of streams and rivers is often assessed with measures of the chemical or physical properties of water. However, a more comprehensive perspective is obtained if resident biological communities are also assessed. Guidelines to protect human health and aquatic life have been established for specific physical and chemical properties of water and have become useful yardsticks with which to assess water quality. Biological communities provide additional crucial information because they live within streams for weeks to years and therefore integrate through time the effects of changes to their chemical or physical environment. In addition, biological communities are a direct measure of stream health—an indicator of the ability of a stream to support aquatic life. Thus, the condition of biological communities, integrated with key physical and chemical properties, provides a comprehensive assessment of stream health.

conterminous United States

Empirical evaluation of the conceptual model underpinning a regional aquatic long-term monitoring program using causal modelling

Conceptual models are an integral facet of long-term monitoring programs. Proposed linkages between drivers, stressors, and ecological indicators are identified within the conceptual model of most mandated programs. We empirically evaluate a conceptual model developed for a regional aquatic and riparian monitoring program using causal models (i.e., Bayesian path analysis). We assess whether data gathered for regional status and trend estimation can also provide insights on why a stream may deviate from reference conditions. We target the hypothesized causal pathways for how anthropogenic drivers of road density, percent grazing, and percent forest within a catchment affect instream biological condition. We found instream temperature and fine sediments in arid sites and only fine sediments in mesic sites accounted for a significant portion of the maximum possible variation explainable in biological condition among managed sites. However, the biological significance of the direct effects of anthropogenic drivers on instream temperature and fine sediments were minimal or not detected. Consequently, there was weak to no biological support for causal pathways related to anthropogenic drivers’ impact on biological condition. With weak biological and statistical effect sizes, ignoring environmental contextual variables and covariates that explain natural heterogeneity would have resulted in no evidence of human impacts on biological integrity in some instances. For programs targeting the effects of anthropogenic activities, it is imperative to identify both land use practices and mechanisms that have led to degraded conditions (i.e., moving beyond simple status and trend estimation). Our empirical evaluation of the conceptual model underpinning the long-term monitoring program provided an opportunity for learning and, consequently, we discuss survey design elements that require modification to achieve question driven monitoring, a necessary step in the practice of adaptive monitoring. We suspect our situation is not unique and many programs may suffer from the same inferential disconnect. Commonly, the survey design is optimized for robust estimates of regional status and trend detection and not necessarily to provide statistical inferences on the causal mechanisms outlined in the conceptual model, even though these relationships are typically used to justify and promote the long-term monitoring of a chosen ecological indicator. Our application demonstrates a process for empirical evaluation of conceptual models and exemplifies the need for such interim assessments in order for programs to evolve and persist.

California, Colorado, Idaho, Montana, Nevada, Oreg

Acidic deposition in the northeastern United States: Sources and inputs, ecosystem effects, and management strategies: The effects of acidic deposition in the northeastern United States include the acidification of soil and water, which stresses terrestrial and aquatic biota

Acidic deposition is the transfer of strong acids and acid-forming substances from the atmosphere to the surface of the Earth. The composition of acidic deposition includes ions, gases, and particles derived from the following: gaseous emissions of sulfur dioxide (SO 2 ), nitrogen oxides (NO x ), ammonia (NH 3 ), and particulate emissions of acidifying and neutralizing compounds. Over the past quarter century of study, acidic deposition has emerged as a critical environmental stress that affects forested landscapes and aquatic ecosystems in North America, Europe, and Asia. This complex problem is an example of a new class of environmental issues that are multiregional in scale and are not amenable to simple resolution by policymakers. Acidic deposition can originate from transboundary air pollution and can affect large geographic areas. It is highly variable across space and time, links air pollution to diverse terrestrial and aquatic ecosystems, and alters the interactions of many elements (e.g., sulfur [S], nitrogen [N], hydrogen ion [H + ], calcium [Ca 2+ ], magnesium [Mg 2+ ], and aluminum [Al]). It also contributes directly and indirectly to biological stress and to the degradation of ecosystems. Despite the complexity of the effects of acidic deposition, North American and European management actions directed toward the recovery of damaged natural resources have resulted in recent decreases in both emissions and deposition of acidic S compounds. Accordingly, acidic deposition presents an instructive case study for coordinating science and policy efforts aimed at resolving large-scale environmental problems.

northeastern United States

Ecological assessment of streams in the Powder River Structural Basin, Wyoming and Montana, 2005-06

Energy and mineral development, particularly coalbed natural gas development, is proceeding at a rapid pace in the Powder River Structural Basin (PRB) in northeastern Wyoming. Concerns about the potential effects of development led to formation of an interagency working group of primarily Federal and State agencies to address these issues in the PRB in Wyoming and in Montana where similar types of resources exist but are largely undeveloped. Under the direction of the interagency working group, an ecological assessment of streams in the PRB was initiated to determine the current status (2005–06) and to establish a baseline for future monitoring. The ecological assessment components include assessment of stream habitat and riparian zones as well as assessments of macroinvertebrate, algal, and fish communities. All of the components were sampled at 47 sites in the PRB during 2005. A reduced set of components, consisting primarily of macroinvertebrate and fish community assessments, was sampled in 2006. Related ecological data, such as habitat and fish community data collected from selected sites in 2004, also are included in this report. The stream habitat assessment included measurement of channel features, substrate size and embeddedness, riparian vegetation, and reachwide characteristics. The width-to-depth ratio (bankfull width/bankfull depth) tended to be higher at sites on the main-stem Powder River than at sites on the main-stem Tongue River and at sites on tributary streams. The streambed substrate particle size was largest at sites on the main-stem Tongue River and smallest at sites on small tributary streams such as Squirrel Creek and Otter Creek. Total vegetative cover at the ground level, understory, and canopy layers ranged from less than 40 percent at a few sites to more than 90 percent at many of the sites. A bank-stability index indicated that sites in the Tongue River drainage were less at risk of bank failure than sites on the main-stem Powder River. Macroinvertebrate communities showed similarity at the river-drainage scale. Macroinvertebrate communities at sites with mountainous headwaters and snowmelt-driven hydrology, such as Clear Creek, Crazy Woman Creek, and Goose Creek, showed similarity with communities from the main-stem Tongue River. The data also indicated similarity among sites on the main-stem Powder River and among small tributaries of the Tongue River. Data analyses using macroinvertebrate observed/expected models and multimetric indices developed by the States of Wyoming and Montana indicated a tendency toward declining biological condition in the downstream direction along the Tongue River. Biological condition for the main-stem Powder River generally improved downstream, from below Salt Creek to near the Wyoming/Montana border, followed by a general decline downstream from the border to the confluence with the Yellowstone River. The biological condition generally was not significantly different between 2005 and 2006, although streamflow was less in 2006 because of drought. Algal communities showed similarity at the river-drainage scale with slight differences from the pattern observed in the macroinvertebrate communities. Although the algal communities from Clear Creek and Goose Creek were similar to those from the main-stem Tongue River, as was true of the macroinvertebrate communities, the algal communities from Crazy Woman Creek had more similarity to those of main-stem Powder River sites than to the Tongue River sites, contrary to the macroinvertebrates. Ordination of algal communities, as well as diatom metrics including salinity and dominant taxa, indicated substantial variation at two sites along the main stem of the Powder River. Fish communities of the PRB were most diverse in the Tongue River drainage. In part due to the effects of Tongue River Reservoir, 15 species of fish were found in the Tongue River drainage that were not found in the Cheyenne, Belle Fourche, or Little Powder River drainages. The number of introduced species and relative abundance of introduced species of fish were higher in the Tongue River and other drainages than at sites on the main-stem Powder River. Although non-native species were identified in the Powder River, the native fish community is largely intact. Western silvery minnow and sturgeon chub—species of special concern—were identified only at sites on the main-stem Powder River and were most common in the Montana segment of the main stem. Fish and habitat sampling on the main-stem Powder River indicated affinity of some species for certain habitats such as pools, runs, riffles, backwaters, or shoals.

Montana, Wyoming

Effects of low pH on the coral reef cryptic invertebrate communities near CO2 vents in Papua New Guinea

Small cryptic invertebrates (the cryptofauna) are extremely abundant, ecologically important, and species rich on coral reefs. Ongoing ocean acidification is likely to have both direct effects on the biology of these organisms, as well as indirect effects through cascading impacts on their habitats and trophic relationships. Naturally acidified habitats have been important model systems for studying these complex interactions because entire communities that are adapted to these environmental conditions can be analyzed. However, few studies have examined the cryptofauna because they are difficult to census quantitatively in topographically complex habitats and are challenging to identify. We addressed these challenges by using Autonomous Reef Monitoring Structures (ARMS) for sampling reef-dwelling invertebrates >2 mm in size and by using DNA barcoding for taxonomic identifications. The study took place in Papua New Guinea at two reef localities, each with three sites at varying distances from carbon dioxide seeps, thereby sampling across a natural gradient in acidification. We observed sharp overall declines in both the abundance (34–56%) and diversity (42–45%) of organisms in ARMS under the lowest pH conditions sampled (7.64–7.75). However, the overall abundance of gastropods increased slightly in lower pH conditions, and crustacean and gastropod families exhibited varying patterns. There was also variability in response between the two localities, despite their close proximity, as one control pH site displayed unusually low diversity and abundances for all invertebrate groups. The data illustrate the complexity of responses of the reef fauna to pH conditions, and the role of additional factors that influence the diversity and abundance of cryptic reef invertebrates.

Milne Bay Province

Irrigation-induced contamination of water, sediment, and biota in the western United States-synthesis of data from the National Irrigation Water Quality Program

In October 1985 the U.S. Department of the Interior (DOI), through the National Irrigation Water Quality Program (NIWQP), began a series of field investigations at 26 areas in the Western United States to determine whether irrigation drainage has had harmful effects on fish, wildlife, and humans or has reduced beneficial uses of water. In 1992 NIWQP initiated the Data Synthesis Project to evaluate data collected during the field investigations. Geologic, climatologic, and hydrologic data were evaluated and water, sediment, and biota from the 26 areas were analyzed to identify commonalities and dominant factors that result in irrigation-induced contamination of water and biota. Data collected for the 26 area investigations have been compiled and merged into a common data base. The structure of the data base is designed to enable assessment of relations between contaminant concentrations in water, sediment, and biota. The data base is available to the scientific community through the World Wide Web at URL http://www.usbr.gov/niwqp. Analysis of the data base for the Data Synthesis included use of summary statistics, factor analysis, and logistic regression. A Geographic Information System was used to store and analyze spatially oriented digital data such as land use, geology and evaporation rates. In the U.S. Department of the Interior (DOI) study areas, samples of water, bottom sediment, and biota were collected for trace-element and pesticide analysis. Contaminants most commonly associated with irrigation drainage were identified by comparing concentrations in water with established criteria. For surface water, the criteria used were typically chronic criteria for the protection of freshwater aquatic life. Because ground water can discharge to the surface where wildlife can be exposed to it, the criteria used for ground water were both the maximum contaminant levels (MCL's) for drinking water and the chronic criteria for the protection of freshwater aquatic life. Data collected by the NIWQP studies indicated that, in surface water, filtered and unfiltered samples had nearly the same concentrations of arsenic, boron, molybdenum, and selenium for concentrations greater than about 10 micrograms per liter. Therefore, in this concentration range, filtered concentrations can be directly compared to biological-effect levels developed for unfiltered samples. In the range of 1 to 10 micrograms per liter there may be a tendency for unfiltered arsenic concentrations to be greater than filtered concentrations. For selenium, however, the data suggest differences from equality in that range result from analytical imprecision and not a general tendency for unfiltered concentrations to be greater than filtered concentrations. This relation may not be true in lentic, nutrient-rich waters because in such settings algae can bioaccumulate large amounts of selenium and other trace elements. Selenium was the trace element in surface water that most commonly exceeded chronic criteria for the protection of freshwater aquatic life; more than 40 percent of the selenium concentrations in surface-water samples exceeded the U.S. Environmental Protection Agency (USEPA) aquatic-life chronic criterion (5 micrograms per liter). In 12 of the 26 areas at least 25 percent of the surface water-samples had selenium concentrations that either equaled or exceeded the chronic criterion (5 micrograms per liter). More than 28 percent of boron concentrations and almost 17 percent of the molybdenum concentrations exceeded the aquatic life criteria established by the State of California (550 and 19 micrograms per liter, respectively). In ground water, more than 22 percent of the arsenic concentrations and more than 35 percent of the selenium concentrations exceeded the MCL (10 and 50 micrograms per liter, respectively). Few samples of uranium in surface water exceeded a criterion for the protection of aquatic life (300 micrograms per liter), but 44 percent

Professional Paper

Extending ordinal regression with a latent zero-augmented beta distribution

Ecological abundance data are often recorded on an ordinal scale in which the lowest category represents species absence. One common example is when plant species cover is visually assessedwithin bounded quadrats and then assigned to pre-defined cover class categories.We present an ordinal beta hurdle model that directly models ordinal category probabilitieswith a biologically realistic beta-distributed latent variable.Ahurdle-at-zero model allows ecologists to explore distribution (absence) and abundance processes in an integrated framework. This provides an alternative to cumulative link models when data are inconsistent with the assumption that the odds ofmoving into a higher category are the same for all categories (proportional odds). Graphical tools and a deviance information criterion were developed to assess whether a hurdle-at-zero model should be used for inferences rather than standard ordinal methods. Hurdle-at-zero and non-hurdle ordinal models fit to vegetation cover class data produced substantially different conclusions.The ordinal beta hurdle model yielded more precise parameter estimates than cumulative logit models, although out-of-sample predictions were similar. The ordinal beta hurdle model provides inferences directly on the latent biological variable of interest, percent cover, and supports exploration of more realistic ecological patterns and processes through the hurdle-at-zero or two-part specification.We provide JAGS code as an on-line supplement. Supplementary materials accompanying this paper appear on-line.

Journal of Agricultural, Biological, and Environme

Assessment of potential effects of water produced from coalbed natural gas development on macroinvertebrate and algal communities in the Powder River and Tongue River, Wyoming and Montana, 2010

Ongoing development of coalbed natural gas in the Powder River structural basin in Wyoming and Montana led to formation of an interagency aquatic task group to address concerns about the effects of the resulting production water on biological communities in streams of the area. Ecological assessments, made from 2005–08 under the direction of the task group, indicated biological condition of the macroinvertebrate and algal communities in the middle reaches of the Powder was lower than in the upper or lower reaches. On the basis of the 2005–08 results, sampling of the macroinvertebrate and algae communities was conducted at 18 sites on the mainstem Powder River and 6 sites on the mainstem Tongue River in 2010. Sampling-site locations were selected on a paired approach, with sites located upstream and downstream of discharge points and tributaries associated with coalbed natural gas development. Differences in biological condition among site pairs were evaluated graphically and statistically using multiple lines of evidence that included macroinvertebrate and algal community metrics (such as taxa richness, relative abundance, functional feeding groups, and tolerance) and output from observed/expected (O/E) macroinvertebrate models from Wyoming and Montana. Multiple lines of evidence indicated a decline in biological condition in the middle reaches of the Powder River, potentially indicating cumulative effects from coalbed natural gas discharges within one or more reaches between Flying E Creek and Wild Horse Creek in Wyoming. The maximum concentrations of alkalinity in the Powder River also occurred in the middle reaches. Biological condition in the upper and lower reaches of the Powder River was variable, with declines between some site pairs, such as upstream and downstream of Dry Fork and Willow Creek, and increases at others, such as upstream and downstream of Beaver Creek. Biological condition at site pairs on the Tongue River showed an increase in one case, near the Wyoming-Montana border, and a decrease in another case, upstream of Tongue River Reservoir. Few significant differences were noted from upstream to downstream of Prairie Dog Creek, a major tributary to the Tongue River. Further study would be needed to confirm the observed patterns and choose areas to examine in greater detail.

Wyoming;Montana

Chemical mixtures and environmental effects: a pilot study to assess ecological exposure and effects in streams

Assessment and management of the risks of exposure to complex chemical mixtures in streams are priorities for human and environmental health organizations around the world. The current lack of information on the composition and variability of environmental mixtures and a limited understanding of their combined effects are fundamental obstacles to timely identification and prevention of adverse human and ecological effects of exposure. This report describes the design of a field-based study of the composition and biological activity of chemical mixtures in U.S. stream waters affected by a wide range of human activities and contaminant sources. The study is a collaborative effort by the U.S. Geological Survey and the U.S. Environmental Protection Agency. Scientists sampled 38 streams spanning 24 States and Puerto Rico. Thirty-four of the sites were located in watersheds impacted by multiple contaminant sources, including industrial and municipal wastewater discharges, crop and animal agricultural runoff, urban runoff, and other point and nonpoint contaminant sources. The remaining four sites were minimally development reference watersheds. All samples underwent comprehensive chemical and biological characterization, including sensitive and specific direct analysis for over 700 dissolved organic and inorganic chemicals and field parameters, identification of unknown contaminants (environmental diagnostics), and a variety of bioassays to evaluate biological activity and toxicity.

Open-File Report

Managing the threat of infectious disease in fisheries and aquaculture using structured decision making

Fisheries and aquaculture provide food and economic security, especially in the developing world, but both face challenges from infectious disease. Here, we consider management of disease issues from a structured decision-making perspective to examine how infectious disease can threaten seafood production and influence management decisions. For both wild fisheries and aquaculture, disease-management objectives generally aim to mitigate the severity and economic burden of outbreaks. General management strategies include manipulating host densities, reducing system connectivity, conserving or improving habitat, and implementing direct treatments or some other biological interventions. To inform decisions, mathematical models can be used to explore disease dynamics and to forecast the potential effectiveness of alternative management actions. Developing and implementing disease-management strategies also involve considering uncertainties and balancing competing stakeholder interests and risk tolerances. We conclude by outlining several steps for applying structured decision making that are broadly useful to decision makers facing issues related to disease.

Frontiers in Ecology and the Environment

Pesticides in small volume plasma samples: Method development and application to smallmouth bass (Micropterus dolomieu) from the Chesapeake Bay watershed, USA

Nontarget organisms are exposed to pesticides following applications in agricultural and urban settings, potentially resulting in deleterious effects. Direct measurements of pesticides in biological tissues may aid in characterizing exposure, accumulation, and potential toxicity versus analyses in environmental media alone (e.g., water, soil, and air). Plasma represents a nonlethal sampling medium that can be used to assess recent exposures to contaminants. Herein, a method was developed to test the extraction of 210 pesticides and their transformation products in small volume plasma samples (100 μL). Plasma samples were protein precipitated with 0.5 % formic acid in acetonitrile added to the sample (ratio of 3.5:1). Pass-through solid phase extraction was used for sample matrix and lipid removal and samples were analyzed by liquid chromatography and gas chromatography with tandem mass spectrometry. Recoveries of 70.0–129.8 % were achieved for 182 pesticides and degradates across the low (25 ng mL −1 ), medium (100 ng mL −1 ), and high (250 ng mL −1 ) spike levels. Method detection levels ranged 0.4–13.0 ng mL −1 . Following development, the method was applied to smallmouth bass ( Micropterus dolomieu ) plasma samples (n = 10) collected from adults in the Chesapeake Bay watershed. Individual plasma samples resulted in four to seven analytes detected with summed concentrations ranging 16.4–95.0 ng mL −1 . Biological multiresidue pesticide methods help elucidate recent exposures of bioactive compounds to nontarget organisms.

Chesapeake Bay watershed

Chronobiologic approach to aquaculture

Diel and seasonal changes in the environment directly or indirectly control many biological functions. Chronobiology, the study of biological functions in relation to time of day, time of year, lunar phases, and other consistent cues, could have important applications in aquaculture. Physiological conditions that vary seasonally include feeding time, tolerance of fish to stress, the state of gonad development, lipid and water content of muscle tissue, and the dress‐out weight of food fish. The quality of cultured fish may vary with the time of day when fish are fed, seined, graded, treated with therapeutic chemicals, injected with hormones, or physically disturbed. Efficiency of production and quality of aquacultural products might be improved if the activities of fish husbandry were timed to coincide with the biological rhythms of fish.

Transactions of the American Fisheries Society

Sonoran Desert: Fragile Land of Extremes

'Sonoran Desert: Fragile Land of Extremes' shows how biologists with the U.S. Geological Survey work with other scientists in an effort to better understand native plants and animals such as desert tortoises, saguaro cacti, and Gila monsters. Much of the program was shot in and around Saguaro National Park near Tucson, Arizona. Genetic detective work, using DNA, focuses on understanding the lives of tortoises. Studies of saguaros over many decades clarify how these amazing plants reproduce and thrive in the desert. Threats from fire, diseases in tortoises, and a growing human population motivate the scientists. Their work to identify how these organisms live and survive is a crucial step for the sound management of biological resources on public lands. This 28-minute program, USGS Open-File Report 03-305, was shot entirely in high definition video and produced by the USGS Western Ecological Research Center and Southwest Biological Science Center; produced and directed by Stephen Wessells, Western Region Office of Communications.

Open-File Report

Integrating bioavailability approaches into waste rock evaluations

The presence of toxic metals in soils affected by mining, industry, agriculture and urbanization, presents problems to human health, the establishment and maintenance of plant and animal habitats, and the rehabilitation of affected areas. A key to managing these problems is predicting the fraction of metal in a given soil that will be biologically labile, and potentially harmful ('bioavailable'). The molecular form of metals and metalloids, particularly the uncomplexed (free) form, controls their bioavailability and toxicity in solution. One computational approach for determining bioavailability, the biotic ligand model (BLM), takes into account not only metal complexation by ligands in solution, but also competitive binding of hardness cations (Ca 2+,Mg 2+,) and metal ions to biological receptor sites. The more direct approach to assess bioavailability is to explicitly measure the response of an organism to a contaminant. A number of microbial enzyme tests have been developed to assess the impact of pollution in a rapid and procedurally simple way. These different approaches in making bioavailability predictions may have value in setting landuse priorities, remediation goals, and habitat reclamation strategies.

Conference Paper

Directional selection shifts trait distributions of planted species in dryland restoration

The match between species trait values and local abiotic filters can restrict community membership. An often-implicit assumption of this relationship is that abiotic filters select for a single locally optimal strategy, though difficulty in isolating effects of the abiotic environment from those of dispersal limitation and biotic interactions has resulted in few empirical tests of this assumption. Similar constraints have made it difficult to assess whether the type and intensity of abiotic filters shift along gradients of environmental harshness, as predicted by the stress-dominance hypothesis. We planted 9,216 plants of 29 perennial grass and forb species that had a range of functional trait values and were assigned to a warm, intermediate or cool temperature tolerance pool across eight sites on the Colorado Plateau. We compared the distributions of traits of surviving individuals to null distributions to evaluate whether there were shifts in trait means and variation. Borrowing from phenotypic selection concepts in evolutionary biology, we assessed support for stabilizing, directional and disruptive abiotic filtering of trait distributions and whether these types of filtering varied with initial species pool. Functional composition was significantly different from null distributions for nearly all traits at all sites, with trait variation more restricted in harsher abiotic conditions, supporting the stress-dominance hypothesis. Contrary to expectations, we primarily found evidence for directional selection, which increased in frequency in warm species pools while disruptive selection was found more often in cool and intermediate species pools. Synthesis . This study provides a controlled experimental approach to test the effect of the abiotic environment on plant trait filtering. We found that opportunistic strategies allowing for rapid water acquisition during favourable periods improved survival at warmer sites. Species with these strategies may be expected to benefit from increasing aridity and may be selected for active management efforts. More generally, the prevalence of directional selection may have important implications for dynamic vegetation models that rely on trait distributions for translating environmental variation into ecosystem processes.

Journal of Ecology

Using indirect methods to constrain symbiotic nitrogen fixation rates: A case study from an Amazonian rain forest

Human activities have profoundly altered the global nitrogen (N) cycle. Increases in anthropogenic N have had multiple effects on the atmosphere, on terrestrial, freshwater and marine ecosystems, and even on human health. Unfortunately, methodological limitations challenge our ability to directly measure natural N inputs via biological N fixation (BNF)-the largest natural source of new N to ecosystems. This confounds efforts to quantify the extent of anthropogenic perturbation to the N cycle. To address this gap, we used a pair of indirect methods -analytical modeling and N balance-to generate independent estimates of BNF in a presumed hotspot of N fixation , a tropical rain forest site in central Rondônia in the Brazilian Amazon Basin. Our objectives were to attempt to constrain symbiotic N fixation rates in this site using indirect methods , and to assess strengths and weaknesses of this approach by looking for areas of convergence and disagreement between the estimates. This approach yielded two remarkably similar estimates of N fixation . However, when compared to a previously published bottom-up estimate, our analysis indicated much lower N inputs via symbiotic BNF in the Rondônia site than has been suggested for the tropics as a whole. This discrepancy may reflect errors associated with extrapolating bottom-up fluxes from plot-scale measures, those resulting from the indirect analyses, and/or the relatively low abundance of legumes at the Rondônia site. While indirect methods have some limitations, we suggest that until the technological challenges of directly measuring N fixation are overcome, integrated approaches that employ a combination of model-generated and empirically-derived data offer a promising way of constraining N inputs via BNF in natural ecosystems.

Biogeochemistry

Pre‐fire drought and competition mediate post‐fire conifer mortality in western U.S. National Parks

Tree mortality is an important outcome of many forest fires. Extensive tree injuries from fire may lead directly to mortality, but environmental and biological stressors may also contribute to tree death. However, there is little evidence showing how the combined effects of two common stressors, drought and competition, influence post‐fire mortality. Geographically broad observations of three common western coniferous trees subjected to prescribed fire showed the likelihood of post‐fire mortality was related to intermediate‐term (10 yr) pre‐fire average radial growth, an important component of tree vigor. Path analysis showed that indices of competition and drought stress prior to fire can be described in terms of joint effects on growth, indirectly affecting post‐fire mortality. Our results suggest that the conditions that govern the relationship between growth and mortality in unburned stands may also apply to post‐fire environments. Thus, biotic and abiotic changes that affect growth negatively (e.g., drought stress) or positively (e.g., growth releases following thinning treatments) prior to fire may influence expressed fire severity, independent of fire intensity (e.g., heat flux, residence time). These relationships suggest that tree mortality may increase under stressful climatic or stand conditions even if fire behavior remains constant.

Ecological Applications