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Showing papers in "Ecological Applications in 2009"


Journal ArticleDOI
TL;DR: It is argued that increased awareness of the implications of spatial bias in surveys, and possible modeling remedies, will substantially improve predictions of species distributions and as large an effect on predictive performance as the choice of modeling method.
Abstract: Most methods for modeling species distributions from occurrence records require additional data representing the range of environmental conditions in the modeled region. These data, called background or pseudo-absence data, are usually drawn at random from the entire region, whereas occurrence collection is often spatially biased toward easily accessed areas. Since the spatial bias generally results in environmental bias, the difference between occurrence collection and background sampling may lead to inaccurate models. To correct the estimation, we propose choosing background data with the same bias as occurrence data. We investigate theoretical and practical implications of this approach. Accurate information about spatial bias is usually lacking, so explicit biased sampling of background sites may not be possible. However, it is likely that an entire target group of species observed by similar methods will share similar bias. We therefore explore the use of all occurrences within a target group as biased background data. We compare model performance using target-group background and randomly sampled background on a comprehensive collection of data for 226 species from diverse regions of the world. We find that target-group background improves average performance for all the modeling methods we consider, with the choice of background data having as large an effect on predictive performance as the choice of modeling method. The performance improvement due to target-group background is greatest when there is strong bias in the target-group presence records. Our approach applies to regression-based modeling methods that have been adapted for use with occurrence data, such as generalized linear or additive models and boosted regression trees, and to Maxent, a probability density estimation method. We argue that increased awareness of the implications of spatial bias in surveys, and possible modeling remedies, will substantially improve predictions of species distributions.

2,307 citations


Journal ArticleDOI
TL;DR: The authors showed that wildfire area burned (WFAB) in the American West was controlled by climate during the 20th century (1916-2003), indicating strong linkages between climate and area burned.
Abstract: The purpose of this paper is to quantify climatic controls on the area burned by fire in different vegetation types in the western United States. We demonstrate that wildfire area burned (WFAB) in the American West was controlled by climate during the 20th century (1916-2003). Persistent ecosystem-specific correlations between climate and WFAB are grouped by vegetation type (ecoprovinces). Most mountainous ecoprovinces exhibit strong year-of-fire relationships with low precipitation, low Palmer drought severity index (PDSI), and high temperature. Grass- and shrub-dominated ecoprovinces had positive relationships with antecedent precipitation or PDSI. For 1977-2003, a few climate variables explain 33-87% (mean = 64%) of WFAB, indicating strong linkages between climate and area burned. For 1916-2003, the relationships are weaker, but climate explained 25-57% (mean = 39%) of the variability. The variance in WFAB is proportional to the mean squared for different data sets at different spatial scales. The importance of antecedent climate (summer drought in forested ecosystems and antecedent winter precipitation in shrub and grassland ecosystems) indicates that the mechanism behind the observed fire-climate relationships is climatic preconditioning of large areas of low fuel moisture via drying of existing fuels or fuel production and drying. The impacts of climate change on fire regimes will therefore vary with the relative energy or water limitations of ecosystems. Ecoprovinces proved a useful compromise between ecologically imprecise state-level and localized gridded fire data. The differences in climate-fire relationships among the ecoprovinces underscore the need to consider ecological context (vegetation, fuels, and seasonal climate) to identify specific climate drivers of WFAB. Despite the possible influence of fire suppression, exclusion, and fuel treatment, WFAB is still substantially controlled by climate. The implications for planning and management are that future WFAB and adaptation to climate change will likely depend on ecosystem-specific, seasonal variation in climate. In fuel-limited ecosystems, fuel treatments can probably mitigate fire vulnerability and increase resilience more readily than in climate-limited ecosystems, in which large severe fires under extreme weather conditions will continue to account for most area burned.

963 citations


Journal ArticleDOI
TL;DR: The thesis is that hierarchical statistical modeling is a powerful way of approaching ecological analysis in the presence of inevitable but quantifiable uncertainties, even if practical issues sometimes require pragmatic compromises.
Abstract: Analyses of ecological data should account for the uncertainty in the process(es) that generated the data. However, accounting for these uncertainties is a difficult task, since ecology is known for its complexity. Measurement and/or process errors are often the only sources of uncertainty modeled when addressing complex ecological problems, yet analyses should also account for uncertainty in sampling design, in model specification, in parameters governing the specified model, and in initial and boundary conditions. Only then can we be confident in the scientific inferences and forecasts made from an analysis. Probability and statistics provide a framework that accounts for multiple sources of uncertainty. Given the complexities of ecological studies, the hierarchical statistical model is an invaluable tool. This approach is not new in ecology, and there are many examples (both Bayesian and non-Bayesian) in the literature illustrating the benefits of this approach. In this article, we provide a baseline for concepts, notation, and methods, from which discussion on hierarchical statistical modeling in ecology can proceed. We have also planted some seeds for discussion and tried to show where the practical difficulties lie. Our thesis is that hierarchical statistical modeling is a powerful way of approaching ecological analysis in the presence of inevitable but quantifiable uncertainties, even if practical issues sometimes require pragmatic compromises.

530 citations


Journal ArticleDOI
TL;DR: This study indicates that natural enemies provide a regionally important ecosystem service by suppressing a key soybean pest, reducing the need for insecticide applications and suggests that management to maintain or enhance landscape diversity has the potential to stabilize or increase biocontrol services.
Abstract: Arthropod predators and parasitoids provide valuable ecosystem services in agricultural crops by suppressing populations of insect herbivores. Many natural enemies are influenced by non-crop habitat surrounding agricultural fields, and understanding if, and at what scales, land use patterns influence natural enemies is essential to predicting how landscape alters biological control services. Here we focus on biological control of soybean aphid, Aphis glycines Matumura, a specialist crop pest recently introduced to the north-central United States. We measured the amount of biological control service supplied to soybean in 26 replicate fields across Michigan, Wisconsin, Iowa, and Minnesota across two years (2005- 2006). We measured the impact of natural enemies by experimentally excluding or allowing access to soybean aphid infested plants and comparing aphid population growth over 14 days. We also monitored aphid and natural enemy populations at large in each field. Predators, principally coccinellid beetles, dominated the natural enemy community of soybean in both years. In the absence of aphid predators, A. glycines increased significantly, with 5.3-fold higher aphid populations on plants in exclusion cages vs. the open field after 14 days. We calculated a biological control services index (BSI) based on relative suppression of aphid populations and related it to landscape diversity and composition at multiple spatial scales surrounding each site. We found that BSI values increased with landscape diversity, measured as Simpson's D. Landscapes dominated by corn and soybean fields provided less biocontrol service to soybean compared with landscapes with an abundance of crop and non-crop habitats. The abundance of Coccinellidae was related to landscape composition, with beetles being more abundant in landscapes with an abundance of forest and grassland compared with landscapes dominated by agricultural crops. Landscape diversity and composition at a scale of 1.5 km surrounding the focal field explained the greatest proportion of the variation in BSI and Coccinellidae abundance. This study indicates that natural enemies provide a regionally important ecosystem service by suppressing a key soybean pest, reducing the need for insecticide applications. Furthermore, it suggests that management to maintain or enhance landscape diversity has the potential to stabilize or increase biocontrol services.

505 citations


Journal ArticleDOI
TL;DR: It is shown how inexpensive, networked digital cameras ("webcams") can be used to document spatial and temporal variation in the spring and autumn phenology of forest canopies, and lays the foundation for regional- to continental-scale camera-based monitoring of phenology at network observatory sites, e.g., National Ecological Observatory Network (NEON) or AmeriFlux.
Abstract: There is a need to document how plant phenology is responding to global change factors, particularly warming trends. "Near-surface" remote sensing, using radiometric instruments or imaging sensors, has great potential to improve phenological monitoring because automated observations can be made at high temporal frequency. Here we build on previous work and show how inexpensive, networked digital cameras ("webcams") can be used to document spatial and temporal variation in the spring and autumn phenology of forest canopies. We use two years of imagery from a deciduous, northern hardwood site, and one year of imagery from a coniferous, boreal transition site. A quantitative signal is obtained by splitting images into separate red, green, and blue color channels and calculating the relative brightness of each channel for "regions of interest" within each image. We put the observed phenological signal in context by relating it to seasonal patterns of gross primary productivity, inferred from eddy covariance measurements of surface-atmosphere CO2 exchange. We show that spring increases, and autumn decreases, in canopy greenness can be detected in both deciduous and coniferous stands. In deciduous stands, an autumn red peak is also observed. The timing and rate of spring development and autumn senescence varies across the canopy, with greater variability in autumn than spring. Interannual variation in phenology can be detected both visually and quantitatively; delayed spring onset in 2007 compared to 2006 is related to a prolonged cold spell from day 85 to day 110. This work lays the foundation for regional- to continental-scale camera-based monitoring of phenology at network observatory sites, e.g., National Ecological Observatory Network (NEON) or AmeriFlux.

444 citations


Journal ArticleDOI
TL;DR: It is suggested that proper site preparation and sustainable harvest practices, such as avoiding the removal or burning of harvest residue, could minimize the impact of afforestation on soils, and slow soil compaction, erosion, and organic matter loss, maintaining soil fertility to the greatest extent possible.
Abstract: Afforestation, the conversion of non-forested lands to forest plantations, can sequester atmospheric carbon dioxide, but the rapid growth and harvesting of biomass may deplete nutrients and degrade soils if managed improperly. The goal of this study is to evaluate how afforestation affects mineral soil quality, including pH, sodium, exchangeable cations, organic carbon, and nitrogen, and to examine the magnitude of these changes regionally where afforestation rates are high. We also examine potential mechanisms to reduce the impacts of afforestation on soils and to maintain long-term productivity. Across diverse plantation types (153 sites) to a depth of 30 cm of mineral soil, we observed significant decreases in nutrient cations (Ca, K, Mg), increases in sodium (Na), or both with afforestation. Across the data set, afforestation reduced soil concentrations of the macronutrient Ca by 29% on average (P , 0.05). Afforestation by Pinus alone decreased soil K by 23% (P , 0.05). Overall, plantations of all genera also led to a mean 71% increase of soil Na (P , 0.05). Mean pH decreased 0.3 units (P , 0.05) with afforestation. Afforestation caused a 6.7% and 15% (P , 0.05) decrease in soil C and N content respectively, though the effect was driven principally by Pinus plantations (15% and 20% decrease, P , 0.05). Carbon to nitrogen ratios in soils under plantations were 5.7-11.6% higher (P , 0.05). In several regions with high rates of afforestation, cumulative losses of N, Ca, and Mg are likely in the range of tens of millions of metric tons. The decreases indicate that trees take up considerable amounts of nutrients from soils; harvesting this biomass repeatedly could impair long-term soil fertility and productivity in some locations. Based on this study and a review of other literature, we suggest that proper site preparation and sustainable harvest practices, such as avoiding the removal or burning of harvest residue, could minimize the impact of afforestation on soils. These sustainable practices would in turn slow soil compaction, erosion, and organic matter loss, maintaining soil fertility to the greatest extent possible.

406 citations


Journal ArticleDOI
TL;DR: Results indicate that mechanical plus fire, fire-only, and mechanical-only treatments using whole-tree harvest systems were all effective at reducing potential fire severity under severe fire weather conditions.
Abstract: Forest structure and species composition in many western U.S. coniferous forests have been altered through fire exclusion, past and ongoing harvesting practices, and livestock grazing over the 20th century. The effects of these activities have been most pronounced in seasonally dry, low and mid-elevation coniferous forests that once experienced frequent, low to moderate intensity, fire regimes. In this paper, we report the effects of Fire and Fire Surrogate (FFS) forest stand treatments on fuel load profiles, potential fire behavior, and fire severity under three weather scenarios from six western U.S. FFS sites. This replicated, multisite experiment provides a framework for drawing broad generalizations about the effectiveness of prescribed fire and mechanical treatments on surface fuel loads, forest structure, and potential fire severity. Mechanical treatments without fire resulted in combined 1-, 10-, and 100-hour surface fuel loads that were significantly greater than controls at three of five FFS sites. Canopy cover was significantly lower than controls at three of five FFS sites with mechanical-only treatments and at all five FFS sites with the mechanical plus burning treatment; fire-only treatments reduced canopy cover at only one site. For the combined treatment of mechanical plus fire, all five FFS sites with this treatment had a substantially lower likelihood of passive crown fire as indicated by the very high torching indices. FFS sites that experienced significant increases in 1-, 10-, and 100-hour combined surface fuel loads utilized harvest systems that left all activity fuels within experimental units. When mechanical treatments were followed by prescribed burning or pile burning, they were the most effective treatment for reducing crown fire potential and predicted tree mortality because of low surface fuel loads and increased vertical and horizontal canopy separation. Results indicate that mechanical plus fire, fire-only, and mechanical-only treatments using whole-tree harvest systems were all effective at reducing potential fire severity under severe fire weather conditions. Retaining the largest trees within stands also increased fire resistance.

367 citations


Journal ArticleDOI
TL;DR: A general framework using the Brownian bridge movement model (BBMM) provides a probabilistic estimate of the migration routes of a sampled population, distinguishes between route segments that function as stopover sites vs. those used primarily as movement corridors, and prioritizes routes for conservation based upon the proportion of the sampled population that uses them.
Abstract: As habitat loss and fragmentation increase across ungulate ranges, identifying and prioritizing migration routes for conservation has taken on new urgency. Here we present a general framework using the Brownian bridge movement model (BBMM) that: (1) provides a probabilistic estimate of the migration routes of a sampled population, (2) distinguishes between route segments that function as stopover sites vs. those used primarily as movement corridors, and (3) prioritizes routes for conservation based upon the proportion of the sampled population that uses them. We applied this approach to a migratory mule deer (Odocoileus hemionus) population in a pristine area of southwest Wyoming, USA, where 2000 gas wells and 1609 km of pipelines and roads have been proposed for development. Our analysis clearly delineated where migration routes occurred relative to proposed development and provided guidance for on-the-ground conservation efforts. Mule deer migration routes were characterized by a series of stopover sites where deer spent most of their time, connected by movement corridors through which deer moved quickly. Our findings suggest management strategies that differentiate between stopover sites and movement corridors may be warranted. Because some migration routes were used by more mule deer than others, proportional level of use may provide a reasonable metric by which routes can be prioritized for conservation. The methods we outline should be applicable to a wide range of species that inhabit regions where migration routes are threatened or poorly understood.

282 citations


Journal ArticleDOI
TL;DR: It is suggested that, while soil resources, including nutrients and moisture, are probably instrumental in determining tree growth rates, disturbances from fire and herbivory may be instrumental in limiting tree cover and facilitating the coexistence of trees and grasses in savannas.
Abstract: Disturbances from fire and herbivory strongly affect savanna vegetation dynamics. In some savannas, fire especially may be instrumental in preserving the coexistence of trees and grasses. The role of herbivory by large mammals is less clear; herbivory has been shown variously to promote and to suppress tree establishment. Here we ask how interactions between herbivory and fire act to shape savanna vegetation dynamics via their effects on tree populations in Hluhluwe iMfolozi Park in KwaZulu Natal, South Africa, a savanna with a full complement of native large mammals. We examined the effects of herbivore exclusion on tree growth, mortality, and seedling establishment from 2000 to 2007 at 10 sites located in areas of low and high herbivore pressure throughout the park. Results were analyzed statistically and using Leslie matrix models of population dynamics. Herbivory and fire acted primarily to suppress sapling growth rather than on sapling mortality or seedling establishment. This indicates that browsing, like fire, suppresses tree density by imposing a demographic bottleneck on the maturation of saplings to adults. Model results suggest that, while browsing and fire each alone impacted growth, a combination of browsing and fire had much greater effects on tree density. Only fire and browsing together were able to prevent increases in tree density. These results suggest that, while soil resources, including nutrients and moisture, are probably instrumental in determining tree growth rates, disturbances from fire and herbivory may be instrumental in limiting tree cover and facilitating the coexistence of trees and grasses in savannas.

275 citations


Journal ArticleDOI
TL;DR: A comprehensive assessment of stable-isotope research on agroecosystem N management can inform the development of policies to mitigate nonpoint source pollution.
Abstract: Intensively managed grain farms are saturated with large inputs of nitrogen (N) fertilizer, leading to N losses and environmental degradation. Despite decades of research directed toward reducing N losses from agroecosystems, progress has been minimal, and the currently promoted best management practices are not necessarily the most effective. We investigated the fate of N additions to temperate grain agroecosystems using a meta-analysis of 217 field-scale studies that followed the stable isotope 15N in crops and soil. We compared management practices that alter inorganic fertilizer additions, such as application timing or reduced N fertilizer rates, to practices that re-couple the biogeochemical cycles of carbon (C) and N, such as organic N sources and diversified crop rotations, and analyzed the following response variables: 15N recovery in crops, total recovery of 15N in crops and soil, and crop yield. More of the literature (94%) emphasized crop recovery of 15N than total 15N recovery in crops and soil (58%), though total recovery is a more ecologically appropriate indicator for assessing N losses. Findings show wide differences in the ability of management practices to improve N use efficiency. Practices that aimed to increase crop uptake of commercial fertilizer had a lower impact on total 15N recovery (3-21% increase) than practices that re-coupled C and N cycling (30-42% increase). A majority of studies (66%) were only one growing season long, which poses a particular problem when organic N sources are used because crops recover N from these sources over several years. These short-term studies neglect significant ecological processes that occur over longer time scales. Field-scale mass balance calculations using the 15N data set show that, on average, 43 kg N x ha(-1) x yr(-1) was unaccounted for at the end of one growing season out of 114 kg N x ha(-1) x yr(-1), representing approximately 38% of the total 15N applied. This comprehensive assessment of stable-isotope research on agroecosystem N management can inform the development of policies to mitigate nonpoint source pollution. Nitrogen management practices that most effectively increase N retention are not currently being promoted and are rare on the landscape in the United States.

255 citations


Journal ArticleDOI
TL;DR: Overall, the response to fuel reduction treatments of the ecological variables presented in this paper was generally maximized by the combined mechanical plus burning treatment.
Abstract: Changes in vegetation and fuels were evaluated from measurements taken before and after fuel reduction treatments (prescribed fire, mechanical treatments, and the combination of the two) at 12 Fire and Fire Surrogate (FFS) sites located in forests with a surface fire regime across the conterminous United States. To test the relative effectiveness of fuel reduction treatments and their effect on ecological parameters we used an information- theoretic approach on a suite of 12 variables representing the overstory (basal area and live tree, sapling, and snag density), the understory (seedling density, shrub cover, and native and alien herbaceous species richness), and the most relevant fuel parameters for wildfire damage (height to live crown, total fuel bed mass, forest floor mass, and woody fuel mass). In the short term (one year after treatment), mechanical treatments were more effective at reducing overstory tree density and basal area and at increasing quadratic mean tree diameter. Prescribed fire treatments were more effective at creating snags, killing seedlings, elevating height to live crown, and reducing surface woody fuels. Overall, the response to fuel reduction treatments of the ecological variables presented in this paper was generally maximized by the combined mechanical plus burning treatment. If the management goal is to quickly produce stands with fewer and larger diameter trees, less surface fuel mass, and greater herbaceous species richness, the combined treatment gave the most desirable results. However, because mechanical plus burning treatments also favored alien species invasion at some sites, monitoring and control need to be part of the prescription when using this treatment.

Journal ArticleDOI
TL;DR: This work develops two new occupancy models for data collected under an increasingly popular sampling design based on spatial replicates that are not selected randomly and that are expected to exhibit Markovian dependence, and fits these models to data from a large-scale tiger occupancy survey recently conducted in Karnataka State, southwestern India.
Abstract: Occupancy modeling focuses on inference about the distribution of organisms over space, using temporal or spatial replication to allow inference about the detection process. Inference based on spatial replication strictly requires that replicates be selected randomly and with replacement, but the importance of these design requirements is not well understood. This paper focuses on an increasingly popular sampling design based on spatial replicates that are not selected randomly and that are expected to exhibit Markovian dependence. We develop two new occupancy models for data collected under this sort of design, one based on an underlying Markov model for spatial dependence and the other based on a trap response model with Markovian detections. We then simulated data under the model for Markovian spatial dependence and fit the data to standard occupancy models and to the two new models. Bias of occupancy estimates was substantial for the standard models, smaller for the new trap response model, and negligible for the new spatial process model. We also fit these models to data from a large-scale tiger occupancy survey recently conducted in Karnataka State, southwestern India. In addition to providing evidence of a positive relationship between tiger occupancy and habitat, model selection statistics and estimates strongly supported the use of the model with Markovian spatial dependence. This new model provides another tool for the decomposition of the detection process, which is sometimes needed for proper estimation and which may also permit interesting biological inferences. In addition to designs employing spatial replication, we note the likely existence of temporal Markovian dependence in many designs using temporal replication. The models developed here will be useful either directly, or with minor extensions, for these designs as well. We believe that these new models represent important additions to the suite of modeling tools now available for occupancy estimation in conservation monitoring. More generally, this work represents a contribution to the topic of cluster sampling for situations in which there is a need for specific modeling (e.g., reflecting dependence) for the distribution of the variable(s) of interest among subunits.

Journal ArticleDOI
TL;DR: A framework for thinking about thresholds in terms of a structured decision making process that focuses directly on the objectives of management, with an aim to providing decisions that are optimal with respect to those objectives.
Abstract: Thresholds and their relevance to conservation have become a major topic of discussion in the ecological literature. Unfortunately, in many cases the lack of a clear conceptual framework for thinking about thresholds may have led to confusion in attempts to apply the concept of thresholds to conservation decisions. Here, we advocate a framework for thinking about thresholds in terms of a structured decision making process. The purpose of this framework is to promote a logical and transparent process for making informed decisions for conservation. Specification of such a framework leads naturally to consideration of definitions and roles of different kinds of thresholds in the process. We distinguish among three categories of thresholds. Ecological thresholds are values of system state variables at which small changes bring about substantial changes in system dynamics. Utility thresholds are components of management objectives (determined by human values) and are values of state or performance variables at which small changes yield substantial changes in the value of the management outcome. Decision thresholds are values of system state variables at which small changes prompt changes in management actions in order to reach specified management objectives. The approach that we present focuses directly on the objectives of management, with an aim to providing decisions that are optimal with respect to those objectives. This approach clearly distinguishes the components of the decision process that are inherently subjective (management objectives, potential management actions) from those that are more objective (system models, estimates of system state). Optimization based on these components then leads to decision matrices specifying optimal actions to be taken at various values of system state variables. Values of state variables separating different actions in such matrices are viewed as decision thresholds. Utility thresholds are included in the objectives component, and ecological thresholds may be embedded in models projecting consequences of management actions. Decision thresholds are determined by the above-listed components of a structured decision process. These components may themselves vary over time, inducing variation in the decision thresholds inherited from them. These dynamic decision thresholds can then be determined using adaptive management. We provide numerical examples (that are based on patch occupancy models) of structured decision processes that include all three kinds of thresholds.

Journal ArticleDOI
TL;DR: This study investigates geographic patterns of coral bleaching in 1998 and 2002 and outlines a synergism between heat stress and nutrient flux as a major causative mechanism for those patterns, providing the first concrete evidence that improved coral reef management will increase the regional-scale survival prospects of coral reefs to global climate change.
Abstract: The threats of wide-scale coral bleaching and reef demise associated with anthropogenic climate change are widely known. Moreover, rates of genetic adaptation and/or changes in the coral-zooxanthella partnerships are considered unlikely to be sufficiently fast for corals to acquire increased physiological resistance to increasing sea temperatures and declining pH. However, it has been suggested that coral reef resilience to climate change may be improved by good local management of coral reefs, including management of water quality. Here, using major data sets from the Great Barrier Reef (GBR), Australia, we investigate geographic patterns of coral bleaching in 1998 and 2002 and outline a synergism between heat stress and nutrient flux as a major causative mechanism for those patterns. The study provides the first concrete evidence for the oft-expressed belief that improved coral reef management will increase the regional-scale survival prospects of coral reefs to global climate change.

Journal ArticleDOI
TL;DR: Estimation of several important components of the carbon balance in forests in Oregon and Northern California during the 1990s found there is still potential to significantly increase the land-based carbon storage by increasing rotation age and reducing harvest rates.
Abstract: Net uptake of carbon from the atmosphere (net ecosystem production, NEP) is dependent on climate, disturbance history, management practices, forest age, and forest type To improve understanding of the influence of these factors on forest carbon stocks and flux in the western United States, federal inventory data and supplemental field measurements at additional plots were used to estimate several important components of the carbon balance in forests in Oregon and Northern California during the 1990s Species- and ecoregion-specific allometric equations were used to estimate live and dead biomass stores, net primary productivity (NPP), and mortality In the semiarid East Cascades and mesic Coast Range, mean total biomass was 8 and 24 kg C/m 2 , and mean NPP was 030 and 078 kg Cm � 2 � yr � 1 , respectively Maximum NPP and dead biomass stores were most influenced by climate, whereas maximum live biomass stores and mortality were most influenced by forest type Within ecoregions, mean live and dead biomass were usually higher on public lands, primarily because of the younger age class distribution on private lands Decrease in NPP with age was not general across ecoregions, with no marked decline in old stands (200 years old) in some ecoregions In the absence of stand-replacing disturbance, total landscape carbon stocks could theoretically increase from 32 6 034 Pg C to 59 6 134 Pg C (a 46% increase) if forests were managed for maximum carbon storage Although the theoretical limit is probably unattainable, given the timber-based economy and fire regimes in some ecoregions, there is still potential to significantly increase the land-based carbon storage by increasing rotation age and reducing harvest rates

Journal ArticleDOI
TL;DR: Through quantifying feedbacks associated with changes in vegetation and those associated withChanges in the snow season length, a more integrated understanding of the manner in which climate change may impact interactions between high-latitude ecosystems and the climate system is reached.
Abstract: 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 (RH) 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 6 0.23 Wm � 2 � 10 yr � 1 (mean 6 SD)), and a positive feedback to climate warming. However, the decrease in albedo due to a shorter snow season (� 5.1 6 1.6 d/10 yr) resulted in much greater regional heat absorption (3.3 6 1.24 Wm � 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.

Journal ArticleDOI
TL;DR: The findings indicate that pines and mesophytic species are sensitive to drought, while oaks are tolerant of drought, and suggest that forest management may be used as a tool to mitigate drought effects.
Abstract: Drought frequency and intensity has been predicted to increase under many climate change scenarios. It is therefore critical to understand the response of forests to potential climate change in an effort to mitigate adverse impacts. The purpose of this study was to explore the regional effects of different drought severities on tree growth and mortality. Specifically, we investigated changes in growth and mortality rates across the southeastern United States under various drought and stand conditions using 1991-2005 Forest Health and Monitoring (FHM) plot data from Alabama, Georgia, and Virginia. Drought effects were examined for three species groups (pines, oaks, and mesophytic species) using the Palmer drought severity index (PDSI) as an indicator of drought severity. Stand variables, including total basal area, total tree density, tree species richness, slope, and stand age, were used to account for drought effects under varying stand conditions. The pines and mesophytic species exhibited significant reductions in growth rate with increasing drought severity. However, no significant difference in growth rate was observed within the oak species group. Mean mortality rates within the no-drought class were significantly lower than those within the other three drought classes, among which no significant differences were found, for both pines and mesophytic species. Mean mortality rates were not significantly different among drought classes for oaks. Total basal area, total tree density, and stand age were negatively related to growth and positively related to mortality, which suggests that older and denser stands are more susceptible to drought damage. The effect of basal area on growth increased with drought severity for the oak and mesophytic species groups. Tree species richness was negatively related to mortality for the pine and mesophytic species groups, indicating that stands with more species suffer less mortality. Slope was positively related to mortality within the mesophytic species group, and its effect increased with drought severity, indicating a higher mortality on sites of greater slope during severe-drought conditions. Our findings indicate that pines and mesophytic species are sensitive to drought, while oaks are tolerant of drought. The observed differential growth and mortality rates among species groups may alter the species composition of southeastern U.S. forests if drought episodes become more frequent and/or intense due to climate change. The significant effects of stand conditions on drought responses observed in our study also suggest that forest management may be used as a tool to mitigate drought effects.

Journal ArticleDOI
TL;DR: This study compares the abilities of volunteer and experienced individuals to detect low-density populations of an actively spreading invasive species, and explores how interobserver variation can bias estimates of the proportion of sites infested derived from occupancy models that allow for both false negative and false positive errors.
Abstract: Monitoring programs increasingly are used to document the spread of invasive species in the hope of detecting and eradicating low-density infestations before they become established. However, interobserver variation in the detection and correct identification of low-density populations of invasive species remains largely unexplored. In this study, we compare the abilities of volunteer and experienced individuals to detect low-density populations of an actively spreading invasive species, and we explore how interobserver variation can bias estimates of the proportion of sites infested derived from occupancy models that allow for both false negative and false positive (misclassification) errors. We found that experienced individuals detected small infestations at sites where volunteers failed to find infestations. However, occupancy models erroneously suggested that experienced observers had a higher probability of falsely detecting the species as present than did volunteers. This unexpected finding is an artifact of the modeling framework and results from a failure of volunteers to detect low-density infestations rather than from false positive errors by experienced observers. Our findings reveal a potential issue with site occupancy models that can arise when volunteer and experienced observers are used together in surveys.

Journal ArticleDOI
TL;DR: Responses of benthic macroinvertebrates along gradients of urban intensity were investigated in nine metropolitan areas across the United States, and threshold analysis showed little evidence for an initial period of resistance to urbanization.
Abstract: Responses of benthic macroinvertebrates along gradients of urban intensity were investigated in nine metropolitan areas across the United States. Invertebrate assemblages in metropolitan areas where forests or shrublands were being converted to urban land were strongly related to urban intensity. In metropolitan areas where agriculture and grazing lands were being converted to urban land, invertebrate assemblages showed much weaker or nonsignificant relations with urban intensity because sites with low urban intensity were already degraded by agriculture. Ordination scores, the number of EPT taxa, and the mean pollution-tolerance value of organisms at a site were the best indicators of changes in assemblage condition. Diversity indices, functional groups, behavior, and dominance metrics were not good indicators of urbanization. Richness metrics were better indicators of urban effects than were abundance metrics, and qualitative samples collected from multiple habitats gave similar results to those of single habitat quantitative samples (riffles or woody snags) in all metropolitan areas. Changes in urban intensity were strongly correlated with a set of landscape variables that was consistent across all metropolitan areas. In contrast, the instream environmental variables that were strongly correlated with urbanization and invertebrate responses varied among metropolitan areas. The natural environmental setting determined the biological, chemical, and physical instream conditions upon which urbanization acts and dictated the differences in responses to urbanization among metropolitan areas. Threshold analysis showed little evidence for an initial period of resistance to urbanization. Instead, assemblages were degraded at very low levels of urbanization, and response rates were either similar across the gradient or higher at low levels of urbanization. Levels of impervious cover that have been suggested as protective of streams (5-10%) were associated with significant assemblage degradation and were not protective.

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TL;DR: The findings indicate that the later autogenic phase strongly influences development trajectories for important wetland soil properties, and the role of different successional phases in determining long-term trajectories of ecosystem development should be considered in restoration design, research, and monitoring.
Abstract: Wetland restoration is increasingly used as a strategy both to address historical wetland losses and to mitigate new wetland impacts. Research has examined the success of restored wetlands for avifaunal habitat, plant biodiversity, and plant cover; however, less is known about soil development in these systems. Soil processes are particularly important as soil organic matter (SOM), cation exchange capacity (CEC), and other properties are directly linked to wetland functions such as water quality improvement. This research compared soil development processes and properties of 30 palustrine depressional wetlands of four different age classes (approximately 5, 14, 35, and 55 years since restoration) located in central New York (USA). Five natural wetlands were used as references. This chronosequence included wetlands 27 years older than previously conducted studies, making it the longest reported database available. Replicated soil cores from each site were analyzed for SOM, bulk density (D(b)), CEC, and concentrations of nutrients and other chemical constituents. Decomposition rate and aboveground plant and litter biomass were measured as key contributors to soil development. The results indicate that some soil properties critical for water quality functions take decades or centuries to reach natural reference levels. Of particular importance, in the top five centimeters of soil, SOM, D(b), and CEC achieved <50% of reference levels 55 years after restoration. Soil development processes in these depressional wetlands appear to be driven by autochthonous inputs and by internal processes such as litter decomposition and are not accelerated in the initial phase of development by allochthonous inputs as has been documented in coastal salt marshes and riverine floodplains. While monitoring generally focuses on the initial establishment phase of restored ecosystems, our findings indicate that the later autogenic phase strongly influences development trajectories for important wetland soil properties. Therefore, the role of different successional phases in determining long-term trajectories of ecosystem development should be considered in restoration design, research, and monitoring. This research highlights areas for improving the field of restoration through understanding of successional processes, increased efforts to jump-start soil development, longer-term monitoring programs, and greater focus on soil components of restored wetlands.

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TL;DR: Grazers can modulate the impact of fire and the strength of the interaction between fire and browsers by altering fuel loads and responding to the distribution of grass across the landscape, and thus exert strong effects on spatial patterns of tree cover in open migratory ecosystems such as the Serengeti.
Abstract: Vertebrate herbivores and fire are known to be important drivers of vegetation dynamics in African savannas. It is of particular importance to understand how changes in herbivore population density, especially of elephants, and fire frequency will affect the amount of tree cover in savanna ecosystems, given the critical importance of tree cover for biodiversity, ecosystem function, and human welfare. We developed a spatially realistic simulation model of vegetation, fire, and dominant herbivore dynamics, tailored to the Serengeti ecosystem of east Africa. The model includes key processes such as tree-grass competition, fire, and resource-based density dependence and adaptive movement by herbivores. We used the model to project the ecosystem 100 years into the future from its present state under different fire, browsing (determined by elephant population density), and grazing (with and without wildebeest present) regimes. The model produced the following key results: (1) elephants and fire exert synergistic negative effects on woody cover; when grazers are excluded, the impact of fire and the strength of the elephant-fire interaction increase; (2) at present population densities of 0.15 elephants/km2, the total amount of woody cover is predicted to remain stable in the absence of fire, but the mature tree population is predicted to decline regardless of the fire regime; without grazers present to mitigate the effects of fire, the size structure of the tree population will become dominated by seedlings and mature trees; (3) spatial heterogeneity in tree cover varies unimodally with elephant population density; fire increases heterogeneity in the presence of grazers and decreases it in their absence; (4) the marked rainfall gradient in the Serengeti directly affects the pattern of tree cover in the absence of fire; with fire, the woody cover is determined by the grazing patterns of the migratory wildebeest, which are partly rainfall driven. Our results show that, in open migratory ecosystems such as the Serengeti, grazers can modulate the impact of fire and the strength of the interaction between fire and browsers by altering fuel loads and responding to the distribution of grass across the landscape, and thus exert strong effects on spatial patterns of tree cover.

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TL;DR: In this article, the authors explored the trade-offs between economic costs, representation targets, and connectivity in identifying spatial priorities across Australia, and found that priority areas that considered ecological and evolutionary processes were more connected along vegetated waterways and were identified for a small increase in economic cost.
Abstract: Systematic conservation planning research has focused on designing systems of conservation areas that efficiently protect a comprehensive and representative set of species and habitats. Recently, there has been an emphasis on improving the adequacy of conservation area design to promote the persistence and future generation of biodiversity. Few studies have explored incorporating ecological and evolutionary processes into conservation planning assessments. Biodiversity in Australia is maintained and generated by numerous ecological and evolutionary processes at various spatial and temporal scales. We accommodated ecological and evolutionary processes in four ways: (1) using sub-catchments as planning units to facilitate the protection of the integrity and function of ecosystem processes occurring on a sub-catchment scale; (2) targeting one type of ecological refugia, drought refugia, which are critical for the persistence of many species during widespread drought; (3) targeting one type of evolutionary refugia which are important for maintaining and generating unique biota during long-term climatic changes; and (4) preferentially grouping priority areas along vegetated waterways to account for the importance of connected waterways and associated riparian areas in maintaining processes. We identified drought refugia, areas of relatively high and regular herbage production in arid and semiarid Australia, from estimates of gross primary productivity derived from satellite data. In this paper, we combined the novel incorporation of these processes with a more traditional framework of efficiently representing a comprehensive sample of biodiversity to identify spatial priorities across Australia. We explored the trade-offs between economic costs, representation targets, and connectivity. Priority areas that considered ecological and evolutionary processes were more connected along vegetated waterways and were identified for a small increase in economic cost. Priority areas for conservation investment are more likely to have long-term benefits to biodiversity if ecological and evolutionary processes are considered in their identification.

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TL;DR: The results presented indicate that otter-trawl impacts are cumulative and can lead to profound changes in benthic communities, which may have far-reaching implications for the integrity of marine food webs and is an important step toward understanding the global ecosystem effects of bottom trawling.
Abstract: Bottom trawling has widespread impacts on benthic communities and habitats. While the direct impacts of trawl disturbances on benthic communities have been extensively studied, the consequences from long-term chronic disturbances are less well understood. The response of benthic macrofauna to chronic otter-trawl disturbance from a Nephrops norvegicus (Norway lobster) fishery was investigated along a gradient of fishing intensity over a muddy fishing ground in the northeastern Irish Sea. Chronic otter trawling had a significant, negative effect on benthic infauna abundance, biomass, and species richness. Benthic epifauna abundance and species richness also showed a significant, negative response, while no such effect was evident for epibenthic biomass. Furthermore, chronic trawl disturbance led to clear changes in community composition of benthic infauna and epifauna. The results presented indicate that otter-trawl impacts are cumulative and can lead to profound changes in benthic communities, which may have far-reaching implications for the integrity of marine food webs. Studies investigating the short-term effects of fishing manipulations previously concluded that otter trawling on muddy substrates had only modest effects on the benthic biota. Hence, the results presented by this study highlight that data from experimental studies can not be readily extrapolated to an ecosystem level and that subtle cumulative effects may only become apparent when fishing disturbances are examined over larger spatial and temporal scales. Furthermore, this study shows that data on chronic effects of bottom trawling on the benthos will be vital in informing the recently advocated move toward an ecosystem approach in fisheries management. As bottom-trawl fisheries are expanding into ever deeper muddy habitats, the results presented here are an important step toward understanding the global ecosystem effects of bottom trawling.

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TL;DR: It is found that the greater the maximum impact caused by an invasive species, the more important it is not only to reduce its density, but also to know the shape of the density-impact relationship accurately.
Abstract: Economic impacts of invasive species worldwide are substantial. Management strategies have been incorporated in population models to assess the effectiveness of management for reducing density, with the implicit assumption that economic impact of the invasive species will also decline. The optimal management effort, however, is that which minimizes the sum of both the management and impact costs. The relationship between population density and economic impact (what we call the “density–impact curve”) is rarely examined in a management context and could take several nonlinear forms. Here we determine the effects of population dynamics and density–impact curves of different shapes on optimal management effort and discover cases where management is either highly effective or a waste of resources. When an inaccurate density–impact curve is used, the increase in total costs due to over- or underinvestment in management can be large. We calculate the increase in total costs incurred if the density–impact curve is incorrect and find that the greater the maximum impact caused by an invasive species, the more important it is not only to reduce its density, but also to know the shape of the density–impact relationship accurately. Lack of information regarding the relationship between density and economic impact causes the most acute problems for invaders that cause high impact at low density, where management typically will be too little, too late. For species that are only problematic at high density, ignorance of the density–impact curve can lead to overinvestment in management with little reduction in impact.

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TL;DR: It is concluded that the severity of this regional drought has masked density-dependent patterns visible in less severe drought conditions and expanded the understanding of stress responses expected in widespread piñon-juniper woodlands.
Abstract: Extreme drought conditions accompanied by rising temperatures have characterized the American Southwest during the past decade, causing widespread tree mortality in pinon-juniper woodlands. Pinon pine (Pinus edulis Engelm.) mortality is linked primarily to outbreaks of the pinyon ips (Ips confusus (Leconte)) precipitated by drought conditions. Although we searched extensively, no biotic agent was identified as responsible for death in Juniperus L. spp. in this study; hence this mortality was due to direct drought stress. Here we examine the relationship between tree abundance and patterns of mortality in three size classes (seedling/sapling, pre-reproductive, reproductive) during the recent extended drought in three regions: southwest Colorado, northern New Mexico, and northern Arizona. Pinon mortality varied from 32% to 65%, and juniper mortality from 3% to 10% across the three sites. In all sites, the greatest pinon mortality was in the larger, presumably older, trees. Using logistic regression models, we examined the influence of tree density and basal area on bark beetle infestations (pinon) and direct drought impacts ( juniper). In contrast to research carried out early in the drought cycle by other researchers in Arizona, we did not find evidence for greater mortality of pinon and juniper trees in increasingly high density or basal area conditions. We conclude that the severity of this regional drought has masked density- dependent patterns visible in less severe drought conditions. With climate projections for the American Southwest suggesting increases in aridity and rising temperatures, it is critical that we expand our understanding of stress responses expected in widespread pinon-juniper woodlands.

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TL;DR: The results suggest that the predicted corridor is robust to uncertainty, and the three carnivore focal species, alone or in combination, were not effective umbrellas for the other focal species.
Abstract: Least-cost models for focal species are widely used to design wildlife corridors. To evaluate the least-cost modeling approach used to develop 15 linkage designs in southern California, USA, we assessed robustness of the largest and least constrained linkage. Species experts parameterized models for eight species with weights for four habitat factors (land cover, topographic position, elevation, road density) and resistance values for each class within a factor (e.g., each class of land cover). Each model produced a proposed corridor for that species. We examined the extent to which uncertainty in factor weights and class resistance values affected two key conservation-relevant outputs, namely, the location and modeled resistance to movement of each proposed corridor. To do so, we compared the proposed corridor to 13 alternative corridors created with parameter sets that spanned the plausible ranges of biological uncertainty in these parameters. Models for five species were highly robust (mean overlap 88%, little or no increase in resistance). Although the proposed corridors for the other three focal species overlapped as little as 0% (mean 58%) of the alternative corridors, resistance in the proposed corridors for these three species was rarely higher than resistance in the alternative corridors (mean difference was 0.025 on a scale of 1- 10; worst difference was 0.39). As long as the model had the correct rank order of resistance values and factor weights, our results suggest that the predicted corridor is robust to uncertainty. The three carnivore focal species, alone or in combination, were not effective umbrellas for the other focal species. The carnivore corridors failed to overlap the predicted corridors of most other focal species and provided relatively high resistance for the other focal species (mean increase of 2.7 resistance units). Least-cost modelers should conduct uncertainty analysis so that decision-makers can appreciate the potential impact of model uncertainty on conservation decisions. Our approach to uncertainty analysis (which can be called a worst-case scenario approach) is appropriate for complex models in which distribution of the input parameters cannot be specified.

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TL;DR: The results provide strong support for regional declines of larger predatory fish in the Baltic Sea promoting algal production by decreasing invertebrate grazer control and highlights the importance of trophic interactions for ecosystem responses to eutrophication.
Abstract: In the Baltic Sea, increased dominance of ephemeral and bloom-forming algae is presently attributed to increased nutrient loads. Simultaneously, coastal predatory fish are in strong decline. Using field data from nine areas covering a 700-km coastline, we examined whether formation of macroalgal blooms could be linked to the composition of the fish community. We then tested whether predator or nutrient availability could explain the field patterns in two small-scale field experiments, by comparing joint effects on algal net production from nutrient enrichment with agricultural fertilizer and exclusion of larger predatory fish with cages. We also manipulated the presence of invertebrate grazers. The abundance of piscivorous fish had a strong negative correlation with the large-scale distribution of bloom-forming macroalgae. Areas with depleted top-predator communities displayed massive increases in their prey, small-bodied fish, and high covers of ephemeral algae. Combining the results from the two experiments showed that excluding larger piscivorous fish: (1) increased the abundance of small-bodied predatory fish; (2) changed the size distribution of the dominating grazers, decreasing the smaller gastropod scrapers; and (3) increased the net production of ephemeral macroalgae. Effects of removing top predators and nutrient enrichment were similar and additive, together increasing the abundance of ephemeral algae many times. Predator effects depended on invertebrate grazers; in the absence of invertebrates there were no significant effects of predator exclusion on algal production. Our results provide strong support for regional declines of larger predatory fish in the Baltic Sea promoting algal production by decreasing invertebrate grazer control. This highlights the importance of trophic interactions for ecosystem responses to eutrophication. The view emerges that to achieve management goals for water quality we need to consider the interplay between top-down and bottom-up processes in future ecosystem management of marine resources.

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TL;DR: Differences in stream ecosystem conditions between pasture reaches and forested sites, including higher stream temperatures, reduced fruit and seed inputs, and a trend toward increased periphyton abundance, appeared to favor fish species normally found in larger streams and facilitate a native invasion process.
Abstract: Riparian forest buffers may play a critical role in moderating the impacts of deforestation on tropical stream ecosystems, but very few studies have examined the ecological effects of riparian buffers in the tropics. To test the hypothesis that riparian forest buffers can reduce the impacts of deforestation on tropical stream biota, we sampled fish assemblages in lowland headwater streams in southeastern Costa Rica representing three different treatments: (1) forested reference stream reaches, (2) stream reaches adjacent to pasture with a riparian forest buffer averaging at least 15 m in width on each bank, and (3) stream reaches adjacent to pasture without a riparian forest buffer. Land cover upstream from the study reaches was dominated by forest at all of the sites, allowing us to isolate the reach-scale effects of the three study treatments. Fish density was significantly higher in pasture reaches than in forest and forest buffer reaches, mostly due to an increase in herbivore-detritivores, but fish biomass did not differ among reach types. Fish species richness was also higher in pasture reaches than in forested reference reaches, while forest buffer reaches were intermediate. Overall, the taxonomic and trophic structure of fish assemblages in forest and forest buffer reaches was very similar, while assemblages in pasture reaches were quite distinct. These patterns were persistent across three sampling periods during our 15-month study. Differences in stream ecosystem conditions between pasture reaches and forested sites, including higher stream temperatures, reduced fruit and seed inputs, and a trend toward increased periphyton abundance, appeared to favor fish species normally found in larger streams and facilitate a native invasion process. Forest buffer reaches, in contrast, had stream temperatures and allochthonous inputs more similar to forested streams. Our results illustrate the importance of riparian areas to stream ecosystem integrity in the tropics and provide support for Costa Rican legislation protecting riparian forests.

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TL;DR: Examination of trends in abundance of four pelagic fish species in the upper San Francisco Estuary over 40 years using Bayesian change point models found no selected covariates could explain statistically the post-2000 change points for any species.
Abstract: We examined trends in abundance of four pelagic fish species (delta smelt, longfin smelt, striped bass, and threadfin shad) in the upper San Francisco Estuary, California, USA, over 40 years using Bayesian change point models. Change point models identify times of abrupt or unusual changes in absolute abundance (step changes) or in rates of change in abundance (trend changes). We coupled Bayesian model selection with linear regression splines to identify biotic or abiotic covariates with the strongest associations with abundances of each species. We then refitted change point models conditional on the selected covariates to explore whether those covariates could explain statistical trends or change points in species abundances. We also fitted a multispecies change point model that identified change points common to all species. All models included hierarchical structures to model data uncertainties, including observation errors and missing covariate values. There were step declines in abundances of all four species in the early 2000s, with a likely common decline in 2002. Abiotic variables, including water clarity, position of the 2 per thousand isohaline (X2), and the volume of freshwater exported from the estuary, explained some variation in species' abundances over the time series, but no selected covariates could explain statistically the post-2000 change points for any species.

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TL;DR: When all phases of the crop rotations were evaluated over the long term, OC decay rates increased concomitantly with OC input rates in several systems, and the quantity of belowground OC inputs was the best predictor of long-term soil C storage.
Abstract: A single ecosystem dominates the Midwestern United States, occupying 26 million hectares in five states alone: the corn-soybean agroecosystem (Zea mays L.-Glycine max (L.) Merr.). Nitrogen (N) fertilization could influence the soil carbon (C) balance in this system because the corn phase is fertilized in 97-100% of farms, at an average rate of 135 kg Nha � 1� yr � 1 . We evaluated the impacts on two major processes that determine the soil C balance, the rates of organic-carbon (OC) inputs and decay, at four levels of N fertilization, 0, 90, 180, and 270 kg/ha, in two long-term experimental sites in Mollisols in Iowa, USA. We compared the corn-soybean system with other experimental cropping systems fertilized with N in the corn phases only: continuous corn for grain; corn-corn-oats (Avena sativa L.)-alfalfa (Medicago sativa L.; corn-oats-alfalfa-alfalfa; and continuous soybean. In all systems, we estimated long-term OC inputs and decay rates over all phases of the rotations, based on long- term yield data, harvest indices (HI), and root : shoot data. For corn, we measured these two ratios in the four N treatments in a single year in each site; for other crops we used published ratios. Total OC inputs were calculated as aboveground plus belowground net primary production (NPP) minus harvested yield. For corn, measured total OC inputs increased with N fertilization (P , 0.05, both sites). Belowground NPP, comprising only 6-22% of total corn NPP, was not significantly influenced by N fertilization. When all phases of the crop rotations were evaluated over the long term, OC decay rates increased concomitantly with OC input rates in several systems. Increases in decay rates with N fertilization apparently offset gains in carbon inputs to the soil in such a way that soil C sequestration was virtually nil in 78% of the systems studied, despite up to 48 years of N additions. The quantity of belowground OC inputs was the best predictor of long-term soil C storage. This indicates that, in these systems, in comparison with increased N-fertilizer additions, selection of crops with high belowground NPP is a more effective management practice for increasing soil C sequestration.