Showing papers in "Ecological Applications in 2016"

Nonnative forest insects and pathogens in the United States: Impacts and policy options.

Abstract: We review and synthesize information on invasions of nonnative forest insects and diseases in the United States, including their ecological and economic impacts, pathways of arrival, distribution within the United States, and policy options for reducing future invasions. Nonnative insects have accumulated in United States forests at a rate of ~2.5 per yr over the last 150 yr. Currently the two major pathways of introduction are importation of live plants and wood packing material such as pallets and crates. Introduced insects and diseases occur in forests and cities throughout the United States, and the problem is particularly severe in the Northeast and Upper Midwest. Nonnative forest pests are the only disturbance agent that has effectively eliminated entire tree species or genera from United States forests within decades. The resulting shift in forest structure and species composition alters ecosystem functions such as productivity, nutrient cycling, and wildlife habitat. In urban and suburban areas, loss of trees from streets, yards, and parks affects aesthetics, property values, shading, stormwater runoff, and human health. The economic damage from nonnative pests is not yet fully known, but is likely in the billions of dollars per year, with the majority of this economic burden borne by municipalities and residential property owners. Current policies for preventing introductions are having positive effects but are insufficient to reduce the influx of pests in the face of burgeoning global trade. Options are available to strengthen the defenses against pest arrival and establishment, including measures taken in the exporting country prior to shipment, measures to ensure clean shipments of plants and wood products, inspections at ports of entry, and post-entry measures such as quarantines, surveillance, and eradication programs. Improved data collection procedures for inspections, greater data accessibility, and better reporting would support better evaluation of policy effectiveness. Lack of additional action places the nation, local municipalities, and property owners at high risk of further damaging and costly invasions. Adopting stronger policies to reduce establishments of new forest insects and diseases would shift the major costs of control to the source and alleviate the economic burden now borne by homeowners and municipalities.

Ecosystem structure, function, and composition in rangelands are negatively affected by livestock grazing.

Abstract: Reports of positive or neutral effects of grazing on plant species richness have prompted calls for livestock grazing to be used as a tool for managing land for conservation. Grazing effects, however, are likely to vary among different response variables, types, and intensity of grazing, and across abiotic conditions. We aimed to examine how grazing affects ecosystem structure, function, and composition. We compiled a database of 7615 records reporting an effect of grazing by sheep and cattle on 278 biotic and abiotic response variables for published studies across Australia. Using these data, we derived three ecosystem measures based on structure, function, and composition, which were compared against six contrasts of grazing pressure, ranging from low to heavy, two different herbivores (sheep, cattle), and across three different climatic zones. Grazing reduced structure (by 35%), function (24%), and composition (10%). Structure and function (but not composition) declined more when grazed by sheep and cattle together than sheep alone. Grazing reduced plant biomass (40%), animal richness (15%), and plant and animal abundance, and plant and litter cover (25%), but had no effect on plant richness nor soil function. The negative effects of grazing on plant biomass, plant cover, and soil function were more pronounced in drier environments. Grazing effects on plant and animal richness and composition were constant, or even declined, with increasing aridity. Our study represents a comprehensive continental assessment of the implications of grazing for managing Australian rangelands. Grazing effects were largely negative, even at very low levels of grazing. Overall, our results suggest that livestock grazing in Australia is unlikely to produce positive outcomes for ecosystem structure, function, and composition or even as a blanket conservation tool unless reduction in specific response variables is an explicit management objective.

Demography of an apex predator at the edge of its range: impacts of changing sea ice on polar bears in Hudson Bay.

Abstract: Changes in the abundance and distribution of wildlife populations are common consequences of historic and contemporary climate change. Some Arctic marine mammals, such as the polar bear (Ursus maritimus), may be particularly vulnerable to such changes due to the loss of Arctic sea ice. We evaluated the impacts of environmental variation on demographic rates for the Western Hudson Bay (WH), polar bear subpopulation from 1984 to 2011 using live-recapture and dead-recovery data in a Bayesian implementation of multistate capture–recapture models. We found that survival of female polar bears was related to the annual timing of sea ice break-up and formation. Using estimated vital rates (e.g., survival and reproduction) in matrix projection models, we calculated the growth rate of the WH subpopulation and projected population responses under different environmental scenarios while accounting for parametric uncertainty, temporal variation, and demographic stochasticity. Our analysis suggested a long-term decline in the number of bears from 1185 (95% Bayesian credible interval [BCI] = 993–1411) in 1987 to 806 (95% BCI = 653–984) in 2011. In the last 10 yr of the study, the number of bears appeared stable due to temporary stability in sea ice conditions (mean population growth rate for the period 2001–2010 = 1.02, 95% BCI = 0.98–1.06). Looking forward, we estimated long-term growth rates for the WH subpopulation of ~1.02 (95% BCI = 1.00–1.05) and 0.97 (95% BCI = 0.92–1.01) under hypothetical high and low sea ice conditions, respectively. Our findings support previous evidence for a demographic linkage between sea ice conditions and polar bear population dynamics. Furthermore, we present a robust framework for sensitivity analysis with respect to continued climate change (e.g., to inform scenario planning) and for evaluating the combined effects of climate change and management actions on the status of wildlife populations.

Influences of prior wildfires on vegetation response to subsequent fire in a reburned Southwestern landscape

Abstract: Large and severe wildfires have raised concerns about the future of forested landscapes in the southwestern United States, especially under repeated burning. In 2011, under extreme weather and drought conditions, the Las Conchas fire burned over several previous burns as well as forests not recently exposed to fire. Our purpose was to examine the influences of prior wildfires on plant community composition and structure, subsequent burn severity, and vegetation response. To assess these relationships, we used satellite-derived measures of burn severity and a nonmetric multidimensional scaling of pre- and post- Las Conchas field samples. Earlier burns were associated with shifts from forested sites to open savannas and meadows, oak scrub, and ruderal communities. These non-forested vegetation types exhibited both resistance to subsequent fire, measured by reduced burn severity, and resilience to reburning, measured by vegetation recovery relative to forests not exposed to recent prior fire. Previous shifts toward non-forested states were strongly reinforced by reburning. Ongoing losses of forests and their ecological values confirm the need for restoration interventions. However, given future wildfire and climate projections, there may also be opportunities presented by transformations toward fire-resistant and resilient vegetation types within portions of the landscape.

Addressing potential local adaptation in species distribution models: Implications for conservation under climate change

Abstract: Species distribution models (SDMs) have been criticized for involving assumptions that ignore or categorize many ecologically relevant factors such as dispersal ability and biotic interactions. Another potential source of model error is the assumption that species are ecologically uniform in their climatic tolerances across their range. Typically, SDMs treat a species as a single entity, although populations of many species differ due to local adaptation or other genetic differentiation. Not taking local adaptation into account may lead to incorrect range prediction and therefore misplaced conservation efforts. A constraint is that we often do not know the degree to which populations are locally adapted. Lacking experimental evidence, we still can evaluate niche differentiation within a species' range to promote better conservation decisions. We explore possible conservation implications of making type I or type II errors in this context. For each of two species, we construct three separate Max-Ent models, one considering the species as a single population and two of disjunct populations. Principal component analyses and response curves indicate different climate characteristics in the current environments of the populations. Model projections into future climates indicate minimal overlap between areas predicted to be climatically suitable by the whole species vs. population-based models. We present a workflow for addressing uncertainty surrounding local adaptation in SDM application and illustrate the value of conducting population-based models to compare with whole-species models. These comparisons might result in more cautious management actions when alternative range outcomes are considered.

Scale-dependent effects of landscape composition and configuration on natural enemy diversity, crop herbivory, and yields.

Abstract: (1) Land-use intensification in agricultural landscapes has led to changes in the way habitats and resources are distributed in space. Pests and their natural enemies are influenced by these changes, and by the farming intensity of crop fields. However, it is unknown whether the composition of landscapes (amount and diversity of land cover types) or their configuration (spatial arrangement of cover types) are more important for natural enemy diversity, and how they impact crop damage and yields. In addition, effects of interactions between local farming practices (organic vs. conventional) and landscape variables are unclear. (2) Here, we make use of a data set where landscape composition and configuration were uncorrelated across multiple spatial scales. Natural enemies, crop damage, and yields were sampled in 35 organic and conventional crop fields. Out of seven broad natural enemy taxa, five were positively affected by a complex landscape configuration. In contrast, only carabids were positively affected by the amount of seminatural habitat around fields. Increasing diversity of land cover types had positive effects on some, but negative effects on other taxa. Effect sizes varied among taxa but increased with increasing spatial scale, defined by circular areas of increasing radius around fields. (3) The diversity of aerial, but not of ground-dwelling enemies was higher in fields under organic than conventional management. Interactions of local and landscape variables were important for birds, but not other enemies. Bird richness was higher in organic fields in simple landscapes, but not in landscapes with complex configuration or high land cover diversity. (4) Crop damage decreased with landscape diversity, but increased in conventional fields with complex configuration. Yields increased with both parameters in conventional fields only, and were higher on average in organic compared to conventional fields. Enemy diversity was positively related to crop damage, indicating positive density-dependence of enemies on pests. However, the diversity of aerial enemies was also positively related to yields. (5) Our results suggest that the effectiveness of agrienvironmental schemes for managing natural enemy diversity, crop damage and yields could be enhanced by optimizing the effects of distinct landscape parameters, particularly landscape configuration and diversity, across scales.

Evaluation of downscaled, gridded climate data for the conterminous United States.

Abstract: Weather and climate affect many ecological processes, making spatially continuous yet fine-resolution weather data desirable for ecological research and predictions. Numerous downscaled weather data sets exist, but little attempt has been made to evaluate them systematically. Here we address this shortcoming by focusing on four major questions: (1) How accurate are downscaled, gridded climate data sets in terms of temperature and precipitation estimates? (2) Are there significant regional differences in accuracy among data sets? (3) How accurate are their mean values compared with extremes? (4) Does their accuracy depend on spatial resolution? We compared eight widely used downscaled data sets that provide gridded daily weather data for recent decades across the United States. We found considerable differences among data sets and between downscaled and weather station data. Temperature is represented more accurately than precipitation, and climate averages are more accurate than weather extremes. The data set exhibiting the best agreement with station data varies among ecoregions. Surprisingly, the accuracy of the data sets does not depend on spatial resolution. Although some inherent differences among data sets and weather station data are to be expected, our findings highlight how much different interpolation methods affect downscaled weather data, even for local comparisons with nearby weather stations located inside a grid cell. More broadly, our results highlight the need for careful consideration among different available data sets in terms of which variables they describe best, where they perform best, and their resolution, when selecting a downscaled weather data set for a given ecological application.

Will climate change increase the risk of plant invasions into mountains

Abstract: Mountain ecosystems have been less adversely affected by invasions of non-native plants than most other ecosystems, partially because most invasive plants in the lowlands are limited by climate and cannot grow under harsher high-elevation conditions. However, with ongoing climate change, invasive species may rapidly move upwards and threaten mid-, and then high-elevation mountain ecosystems. We evaluated this threat by modeling the current and future habitat suitability for 48 invasive plant species in Switzerland and New South Wales, Australia. Both regions had contrasting climate interactions with elevation, resulting in possible different responses of species distributions to climate change. Using a species distribution modeling approach that combines data from two spatial scales, we built high-resolution species distribution models (≤ 250 m) that account for the global climatic niche of species and also finer variables depicting local climate and disturbances. We found that different environmental drivers limit the elevation range of invasive species in each of the two regions, leading to region-specific species responses to climate change. The optimal suitability for plant invaders is predicted to markedly shift from the lowland to the montane or subalpine zone in Switzerland, whereas the upward shift is far less pronounced in New South Wales where montane and subalpine elevations are already suitable. The results suggest that species most likely to invade high elevations in Switzerland will be cold-tolerant, whereas species with an affinity to moist soils are most likely to invade higher elevations in Australia. Other plant traits were only marginally associated with elevation limits. These results demonstrate that a more systematic consideration of future distributions of invasive species is required in conservation plans of not yet invaded mountainous ecosystems.

Season and landscape composition affect pollen foraging distances and habitat use of honey bees

Abstract: Honey bees (Apis mellifera L.) show a large variation in foraging distances and use a broad range of plant species as pollen resources, even in regions with intensive agriculture. However, it is unknown how increasing areas of mass-flowering crops like oilseed rape (Brassica napus; OSR) or a decrease of seminatural habitats (SNH) change the temporal and spatial availability of pollen resources for honey bee colonies, and thus foraging distances and frequency in different habitat types. We studied pollen foraging of honey bee colonies in 16 agricultural landscapes with independent gradients of OSR and SNH area within 2 km and used waggle dances and digital geographic maps with major land cover types to reveal the distance and visited habitat type on a landscape level. Mean pollen foraging distance of 1347 decoded bee dances was 1015 m (± 26 m; SEM). In spring, increasing area of flowering OSR within 2 km reduced mean pollen foraging distances from 1324 m to only 435 m. In summer, increasing cover of SNH areas close to the colonies (within 200 m radius) reduced mean pollen foraging distances from 846 to 469 m. Frequency of pollen foragers per habitat type, measured as the number of dances per hour and hectare, was equally high for SNH, grassland, and OSR fields, but lower for other crops and forests. In landscapes with a small proportion of SNH a significantly higher density of pollen foragers on SNH was observed, indicating that pollen resources in such simple agricultural landscapes are more limited. Overall, we conclude that SNH and mass-flowering crops can reduce foraging distances of honey bee colonies at different scales and seasons with possible benefits for the performance of honey bee colonies. Further, mixed agricultural landscapes with a high proportion of SNH reduce foraging densities of honey bees in SNH and thus possible competition for pollen resources.

Characterizing driver-response relationships in marine pelagic ecosystems for improved ocean management.

Abstract: Scientists and resource managers often use methods and tools that assume ecosystem components respond linearly to environmental drivers and human stressors. However, a growing body of literature demonstrates that many relationships are-non-linear, where small changes in a driver prompt a disproportionately large ecological response. We aim to provide a comprehensive assessment of the relationships between drivers and ecosystem components to identify where and when non-linearities are likely to occur. We focused our analyses on one of the best-studied marine systems, pelagic ecosystems, which allowed us to apply robust statistical techniques on a large pool of previously published studies. In this synthesis, we (1) conduct a wide literature review on single driver-response relationships in pelagic systems, (2) use statistical models to identify the degree of non-linearity in these relationships, and (3) assess whether general patterns exist in the strengths and shapes of non-linear relationships across drivers. Overall we found that non-linearities are common in pelagic ecosystems, comprising at least 52% of all driver-response relation- ships. This is likely an underestimate, as papers with higher quality data and analytical approaches reported non-linear relationships at a higher frequency (on average 11% more). Consequently, in the absence of evidence for a linear relationship, it is safer to assume a relationship is non-linear. Strong non-linearities can lead to greater ecological and socioeconomic consequences if they are unknown (and/or unanticipated), but if known they may provide clear thresholds to inform management targets. In pelagic systems, strongly non-linear relationships are often driven by climate and trophodynamic variables but are also associated with local stressors, such as overfishing and pollution, that can be more easily controlled by managers. Even when marine resource managers cannot influence ecosystem change, they can use information about threshold responses to guide how other stressors are managed and to adapt to new ocean conditions. As methods to detect and reduce uncertainty around threshold values improve, managers will be able to better understand and account for ubiquitous non-linear relationships.

Rapidly restoring biological soil crusts and ecosystem functions in a severely disturbed desert ecosystem.

Abstract: Restoring biological soil crusts (biocrusts) in degraded drylands can contribute to recovery of ecosystem functions that have global implications, including erosion resistance and nutrient cycling. To examine techniques for restoring biocrusts, we conducted a replicated, factorial experiment on recently abandoned road surfaces by applying biocrust inoculation (salvaged and stored dry for two years), salvaged topsoil, an abiotic soil amendment (wood shavings), and planting of a dominant perennial shrub (Ambrosia dumosa). Eighteen months after treatments, we measured biocrust abundance and species composition, soil chlorophyll a content and fertility, and soil resistance to erosion. Biocrust addition significantly accelerated biocrust recovery on disturbed soils, including increasing lichen and moss cover and cyanobacteria colonization. Compared to undisturbed controls, inoculated plots had similar lichen and moss composition, recovered 43% of total cyanobacteria density, had similar soil chlorophyll content, and exhibited recovery of soil fertility and soil stability. Inoculation was the only treatment that generated lichen and moss cover. Topsoil application resulted in partial recovery of the cyanobacteria community and soil properties. Compared to untreated disturbed plots, topsoil application without inoculum increased cyanobacteria density by 186% and moderately improved soil chlorophyll and ammonium content and soil stability. Topsoil application produced 22% and 51% of the cyanobacteria density g⁻¹ soil compared to undisturbed and inoculated plots, respectively. Plots not treated with either topsoil or inoculum had significantly lower cyanobacteria density, soil chlorophyll and ammonium concentrations, and significantly higher soil nitrate concentration. Wood shavings and Ambrosia had no influence on biocrust lichen and moss species recovery but did affect cyanobacteria composition and soil fertility. Inoculation of severely disturbed soil with native biocrusts rapidly restored biocrust communities and soil stability such that restored areas were similar to undisturbed desert within three years. Using salvaged biocrust as inoculum can be an effective tool in ecological restoration because of its efficacy and simple implementation. Although salvaging biocrust material can be technically difficult and potentially costly, utilizing opportunities to salvage material in planned future disturbance can provide additional land management tools.

Fortifying the forest: thinning and burning increase resistance to a bark beetle outbreak and promote forest resilience.

Abstract: Fire frequency in low-elevation coniferous forests in western North America has greatly declined since the late 1800s. In many areas, this has increased tree density and the proportion of shade-tolerant species, reduced resource availability, and increased forest susceptibility to forest insect pests and high-severity wildfire. In response, treatments are often implemented with the goal of increasing ecosystem resilience by increasing resistance to disturbance. We capitalized on an existing replicated study of fire and stand density treatments in a ponderosa pine (Pinus ponderosa)-Douglas-fir (Pseudotsuga menziesii) forest in western Montana, U.S. that experienced a naturally occurring mountain pine beetle (MPB; Dendroctonus ponderosae) outbreak five years after implementation of fuels treatments. We explored whether treatment effects on tree-level defense and stand structure affected resistance to MPB. Mortality from MPB was highest in the denser, untreated control and burn-only treatments, with approximately 50% and 39%, respectively, of ponderosa pine killed during the outbreak, compared to almost no mortality in the thin-only and thin-burn treatments. Thinning treatments, with or without fire, dramatically increased tree growth and resin ducts relative to control and burn-only treatments. Prescribed burning did not increase resin ducts, but did cause changes in resin chemistry that may have affected MPB communication and lowered attack success. While ponderosa pine remained dominant in the thin and thin-burn treatments after the outbreak, the high pine mortality in the control and burn-only treatment caused a shift in species dominance to Douglas-fir. The high Douglas-fir component in the control and burn-only treatments due to 20th century fire exclusion, coupled with high pine mortality from MPB, has likely reduced resilience of this forest beyond the ability to return to a ponderosa pine-dominated system in the absence of further fire or mechanical treatment. Our results show treatments designed to increase resistance to high-severity fire in ponderosa pine-dominated forests in the Northern Rockies can also increase resistance to MPB, even during an outbreak. This study suggests that fuel and restoration treatments in fire-dependent ponderosa pine forests that reduce tree density increase ecosystem resilience in the short term, while the reintroduction of fire is important for long-term resilience. This article is protected by copyright. All rights reserved.

Effects of breeding versus winter habitat loss and fragmentation on the population dynamics of a migratory songbird

Abstract: Many migratory species are in decline and understanding these declines is challenging because individuals occupy widely divergent and geographically distant habitats during a single year and therefore populations across the range are interconnected in complex ways. Network modeling has been used to show, theoretically, that shifts in migratory connectivity patterns can occur in response to habitat or climate changes and that habitat loss in one region can affect sub-populations in regions that are not directly connected. Here, we use a network model, parameterized by integrating long-term monitoring data with direct tracking of ~100 individuals, to explain population trends in the rapidly declining wood thrush (Hylocichla mustelina), and to predict future trends. Our model suggests that species-level declines in wood thrush are driven primarily by tropical deforestation in Central America but that protection of breeding habitat in some regions is necessary to prevent shifts in migratory connectivity and t...

Threshold effect of habitat loss on bat richness in cerrado-forest landscapes

Abstract: Understanding how animal groups respond to contemporary habitat loss and fragmentation is essential for development of strategies for species conservation. Until now, there has been no consensus about how landscape degradation affects the diversity and distribution of Neotropical bats. Some studies demonstrate population declines and species loss in impacted areas, although the magnitude and generality of these effects on bat community structure are unclear. Empirical fragmentation thresholds predict an accentuated drop in biodiversity, and species richness in particular, when less than 30% of the original amount of habitat in the landscape remains. In this study, we tested whether bat species richness demonstrates this threshold response, based on 48 sites distributed across 12 landscapes with 9-88% remaining forest in Brazilian cerrado-forest formations. We also examined the degree to which abundance was similarly affected within four different feeding guilds. The threshold value for richness, below which bat diversity declines precipitously, was estimated at 47% of remaining forest. To verify if the response of bat abundance to habitat loss differed among feeding guilds, we used a model selection approach based on Akaike's information criterion. Models accounted for the amount of riparian forest, semideciduous forest, cerrado, tree plantations, secondary forest, and the total amount of forest in the landscape. We demonstrate a nonlinear effect of the contribution of tree plantations to frugivores, and a positive effect of the amount of cerrado to nectarivores and animalivores, the groups that responded most to decreases in amount of forest. We suggest that bat assemblages in interior Atlantic Forest and cerrado regions of southeastern Brazil are impoverished, since we found lower richness and abundance of different groups in landscapes with lower amounts of forest. The relatively higher threshold value of 47% suggests that bat communities have a relatively lower resistance to habitat degradation than other animal groups. Accordingly, conservation and restoration strategies should focus on increasing the amount of native vegetation of landscapes so as to enhance species richness of bats.

Protected areas and their surrounding territory: socioecological systems in the context of ecological solidarity

Abstract: The concept of ecological solidarity (ES) is a major feature of the 2006 law reforming national park policy in France. In the context of biodiversity conservation, the objectives of this study are to outline the historical development of ES, provide a working definition, and present a method for its implementation that combines environmental pragmatism and adaptive management. First, we highlight how ES provides a focus on the interdependencies among humans and nonhuman components of the socioecological system. In doing so, we identify ES within a framework that distinguishes ecological, socioecological, and sociopolitical interdependencies. In making such interdependencies apparent to humans who are not aware of their existence, the concept of ES promotes collective action as an alternative or complementary approach to state- or market-based approaches. By focusing on the awareness, feelings, and acknowledgement of interdependencies between actors and between humans and nonhumans, we present and discuss a learning-based approach (participatory modeling) that allows stakeholders to work together to construct cultural landscapes for present and future generations. Using two case studies, we show how an ES analysis goes beyond the ecosystem management approach to take into account how human interactions with the environment embody cultural, social, and economic values and endorse an ethically integrated science of care and responsibility. ES recognizes the diversity of these values as a practical foundation for socially engaged and accountable actions. Finally, we discuss how ES enhances academic support for a socioecological systems approach to biodiversity conservation and promotes collaboration with decision-makers and stakeholders involved in the adaptive management of protected areas and their surrounding landscapes.

Comparison of solar-induced chlorophyll fluorescence, light-use efficiency, and process-based GPP models in maize

Abstract: Accurately quantifying cropland gross primary production (GPP) is of great importance to monitor cropland status and carbon budgets. Satellite-based light-use efficiency (LUE) models and process-based terrestrial biosphere models (TBMs) have been widely used to quantify cropland GPP at different scales in past decades. However, model estimates of GPP are still subject to large uncertainties, especially for croplands. More recently, space-borne solar-induced chlorophyll fluorescence (SIF) has shown the ability to monitor photosynthesis from space, providing new insights into actual photosynthesis monitoring. In this study, we examined the potential of SIF data to describe maize phenology and evaluated three GPP modeling approaches (space-borne SIF retrievals, a LUE-based vegetation photosynthesis model [VPM], and a process-based soil canopy observation of photochemistry and energy flux [SCOPE] model constrained by SIF) at a maize (Zea mays L.) site in Mead, Nebraska, USA. The result shows that SIF captured the seasonal variations (particularly during the early and late growing season) of tower-derived GPP (GPP_EC) much better than did satellite-based vegetation indices (enhanced vegetation index [EVI] and land surface water index [LSWI]). Consequently, SIF was strongly correlated with GPP_EC than were EVI and LSWI. Evaluation of GPP estimates against GPP_EC during the growing season demonstrated that all three modeling approaches provided reasonable estimates of maize GPP, with Pearson's correlation coefficients (r) of 0.97, 0.94, and 0.93 for the SCOPE, VPM, and SIF models, respectively. The SCOPE model provided the best simulation of maize GPP when SIF observations were incorporated through optimizing the key parameter of maximum carboxylation capacity (Vcmax). Our results illustrate the potential of SIF data to offer an additional way to investigate the seasonality of photosynthetic activity, to constrain process-based models for improving GPP estimates, and to reasonably estimate GPP by integrating SIF and GPP_EC data without dependency on climate inputs and satellite-based vegetation indices.

Grazing intensity differentially regulates ANPP response to precipitation in North American semiarid grasslands

Abstract: Grazing intensity elicits changes in the composition of plant functional groups in both shortgrass steppe (SGS) and northern mixed-grass prairie (NMP) in North America. How these grazing intensity-induced changes control aboveground net primary production (ANPP) responses to precipitation remains a central open question, especially in light of predicted climate changes. Here, we evaluated effects of four levels (none, light, moderate, and heavy) of long-term (>30 yr) grazing intensity in SGS and NMP on: (1) ANPP; (2) precipitation-use efficiency (PUE, ANPP : precipitation); and (3) precipitation marginal response (PMR; slope of a linear regression model between ANPP and precipitation). We advance prior work by examining: (1) the consequences of a range of grazing intensities (more grazed vs. ungrazed); and (2) how grazing-induced changes in ANPP and PUE are related both to shifts in functional group composition and physiological responses within each functional group. Spring (April-June) precipitation, the primary determinant of ANPP, was only 12% higher in NMP than in SGS, yet ANPP and PUE were 25% higher. Doubling grazing intensity in SGS and nearly doubling it in NMP reduced ANPP and PUE by only 24% and 33%, respectively. Increased grazing intensity reduced C3 graminoid biomass and increased C4 grass biomass in both grasslands. Functional group shifts affected PUE through biomass reductions, as PUE was positively associated with the relative abundance of C3 species and negatively with C4 species across both grasslands. At the community level, PMR was similar between grasslands and unaffected by grazing intensity. However, PMR of C3 graminoids in SGS was eightfold higher in the ungrazed treatment than under any grazed level. In NMP, PMR of C3 graminoids was only reduced under heavy grazing intensity. Knowing the ecological consequences of grazing intensity provides valuable information for mitigation and adaptation strategies in response to predicted climate change. For example, moderate grazing (the recommended rate) in SGS would sequester the same amount of aboveground carbon as light grazing because ANPP was nearly the same. In contrast, reductions in grazing intensity in NMP from moderate to light intensity would increase the amount of aboveground carbon sequestrated by 25% because of increased ANPP.

Repeated wildfires alter forest recovery of mixed‐conifer ecosystems

Abstract: Most models project warmer and drier climates that will contribute to larger and more frequent wildfires. However, it remains unknown how repeated wildfires alter post-fire successional patterns and forest structure. Here, we test the hypothesis that the number of wildfires, as well as the order and severity of wildfire events interact to alter forest structure and vegetation recovery and implications for vegetation management. In 2014, we examined forest structure, composition, and tree regeneration in stands that burned 1-18 yr before a subsequent 2007 wildfire. Three important findings emerged: (1) Repeatedly burned forests had 15% less woody surface fuels and 31% lower tree seedling densities compared with forests that only experienced one recent wildfire. These repeatedly burned areas are recovering differently than sites burned once, which may lead to alternative ecosystem structure. (2) Order of burn severity (high followed by low severity compared with low followed by high severity) did influence forest characteristics. When low burn severity followed high, forests had 60% lower canopy closure and total basal area with 92% fewer tree seedlings than when high burn severity followed low. (3) Time between fires had no effect on most variables measured following the second fire except large woody fuels, canopy closure and tree seedling density. We conclude that repeatedly burned areas meet many vegetation management objectives of reduced fuel loads and moderate tree seedling densities. These differences in forest structure, composition, and tree regeneration have implications not only for the trajectories of these forests, but may reduce fire intensity and burn severity of subsequent wildfires and may be used in conjunction with future fire suppression tactics.

Heavy and frequent thinning promotes drought adaptation in Pinus sylvestris forests.

Abstract: Droughts and their negative effects on forest ecosystems are projected to increase under climate change for many regions. It has been suggested that intensive thinning could reduce drought impacts on established forests in the short-term. Most previous studies on the effect of thinning on drought impacts, however, have been confined to single forest sites. It is therefore still unclear how general and persisting the benefits of thinning are. This study assesses the potential of thinning to increase drought tolerance of the wide spread Scots pine (Pinus sylvestris) in Central Europe. We hypothesized (1) that increasing thinning intensity benefits the maintenance of radial growth of crop trees during drought (resistance) and its recovery following drought, (2) that those benefits to growth decrease with time elapsed since the last thinning and with stand age, and (3) that they may depend on drought severity as well as water limitations in pre- and post-drought periods. To test these hypotheses, we assessed the effects of thinning regime, stand age, and drought severity on radial growth of 129 Scots pine trees during and after drought events in four long-term thinning experiments in Germany. We found that thinning improved the recovery of radial growth following drought and to a lesser extent the growth resistance during a drought event. Growth recovery following drought was highest after the first thinning intervention and in recently and heavily thinned stands. With time since the last thinning, however, this effect decreased and could even become negative when compared to unthinned stands. Further, thinning helped to avoid an age-related decline in growth resistance (and recovery) following drought. The recovery following drought, but not the resistance during drought, was related to water limitations in the drought period. This is the first study that analyzed drought-related radial growth in trees of one species across several stands of different age. The interaction between thinning intensity and time since the last thinning underline the importance to distinguish between short- and long-term effects of thinning. According to our analysis, only thinning regimes, with relatively heavy and frequent thinning interventions would increase drought tolerance in pine stands.

Disturbance gradient shows logging affects plant functional groups more than fire

Abstract: Understanding the impacts of natural and human disturbances on forest biota is critical for improving forest management. Many studies have examined the separate impacts on fauna and flora of wildfire, conventional logging, and salvage logging, but empirical comparisons across a broad gradient of simultaneous disturbances are lacking. We quantified species richness and frequency of occurrence of vascular plants, and functional group responses, across a gradient of disturbances that occurred concurrently in 2009 in the mountain ash forests of southeastern Australia. Our study encompassed replicated sites in undisturbed forest (~70 yr post fire), forest burned at low severity, forest burned at high severity, unburned forest that was clearcut logged, and forest burned at high severity that was clearcut salvage logged post-fire. All sites were sampled 2 and 3 yr post fire. Mean species richness decreased across the disturbance gradient from 30.1 species/site on low-severity burned sites and 28.9 species/site on high-severity burned sites, to 25.1 species/site on clearcut sites and 21.7 species/site on salvage logged sites. Low-severity burned sites were significantly more species-rich than clearcut sites and salvage logged sites; high-severity burned sites supported greater species richness than salvage logged sites. Specific traits influenced species' sensitivity to disturbance. Resprouting species dominated undisturbed mountain ash forests, but declined significantly across the gradient. Fern and midstory trees decreased significantly in frequency of occurrence across the gradient. Ferns (excluding bracken) decreased from 34% of plants in undisturbed forest to 3% on salvage logged sites. High-severity burned sites supported a greater frequency of occurrence and species richness of midstory trees compared to clearcut and salvage logged sites. Salvage logging supported fewer midstory trees than any other disturbance category, and were distinctly different from clearcut sites. Plant life form groups, including midstory trees, shrubs, and ferns, were dominated by very few species on logged sites. The differences in biotic response across the gradient of natural and human disturbances have significant management implications, particularly the need to reduce mechanical disturbance overall and to leave specific areas with no mechanical disturbance across the cut area during logging operations, to ensure the persistence of resprouting taxa.

A framework for inferring biological communities from environmental DNA

Abstract: Environmental DNA (eDNA), genetic material recovered from an environmental medium such as soil, water, or feces, reflects the membership of the ecological community present in the sampled environment. As such, eDNA is a potentially rich source of data for basic ecology, conservation, and management, because it offers the prospect of quantitatively reconstructing whole ecological communities from easily obtained samples. However, like all sampling methods, eDNA sequencing is subject to methodological limitations that can generate biased descriptions of ecological communities. Here, we demonstrate parallels between eDNA sampling and traditional sampling techniques, and use these parallels to offer a statistical structure for framing the challenges faced by eDNA and for illuminating the gaps in our current knowledge. Although the current state of knowledge on some of these steps precludes a full estimate of biomass for each taxon in a sampled eDNA community, we provide a map that illustrates potential methods for bridging these gaps. Additionally, we use an original data set to estimate the relative abundances of taxon-specific template DNA prior to PCR, given the abundance of DNA sequences recovered post-PCR-and-sequencing, a critical step in the chain of eDNA inference. While we focus on the use of eDNA samples to determine the relative abundance of taxa within a community, our approach also applies to single-taxon applications (including applications using qPCR), studies of diversity, and studies focused on occurrence. By grounding inferences about eDNA community composition in a rigorous statistical framework, and by making these inferences explicit, we hope to improve the inferential potential for the emerging field of community-level eDNA analysis.

Multiscale connectivity and graph theory highlight critical areas for conservation under climate change.

Abstract: Conservation planning and biodiversity management require information on landscape connectivity across a range of spatial scales from individual home ranges to large regions. Reduction in landscape connectivity due changes in land use or development is expected to act synergistically with alterations to habitat mosaic configuration arising from climate change. We illustrate a multiscale connectivity framework to aid habitat conservation prioritization in the context of changing land use and climate. Our approach, which builds upon the strengths of multiple landscape connectivity methods, including graph theory, circuit theory, and least-cost path analysis, is here applied to the conservation planning requirements of the Mohave ground squirrel. The distribution of this threatened Californian species, as for numerous other desert species, overlaps with the proposed placement of several utility-scale renewable energy developments in the American southwest. Our approach uses information derived at three spatial scales to forecast potential changes in habitat connectivity under various scenarios of energy development and climate change. By disentangling the potential effects of habitat loss and fragmentation across multiple scales, we identify priority conservation areas for both core habitat and critical corridor or stepping stone habitats. This approach is a first step toward applying graph theory to analyze habitat connectivity for species with continuously distributed habitat and should be applicable across a broad range of taxa.

Climate influences on whitebark pine mortality from mountain pine beetle in the Greater Yellowstone Ecosystem.

Abstract: Extensive mortality of whitebark pine, beginning in the early to mid-2000s, occurred in the Greater Yellowstone Ecosystem (GYE) of the western USA, primarily from mountain pine beetle but also from other threats such as white pine blister rust. The climatic drivers of this recent mortality and the potential for future whitebark pine mortality from mountain pine beetle are not well understood, yet are important considerations in whether to list whitebark pine as a threatened or endangered species. We sought to increase the understanding of climate influences on mountain pine beetle outbreaks in whitebark pine forests, which are less well understood than in lodgepole pine, by quantifying climate–beetle relationships, analyzing climate influences during the recent outbreak, and estimating the suitability of future climate for beetle outbreaks. We developed a statistical model of the probability of whitebark pine mortality in the GYE that included temperature effects on beetle development and survival, precipitation effects on host tree condition, beetle population size, and stand characteristics. Estimated probability of whitebark pine mortality increased with higher winter minimum temperature, indicating greater beetle winter survival; higher fall temperature, indicating synchronous beetle emergence; lower two-year summer precipitation, indicating increased potential for host tree stress; increasing beetle populations; stand age; and increasing percent composition of whitebark pine within a stand. The recent outbreak occurred during a period of higher-than-normal regional winter temperatures, suitable fall temperatures, and low summer precipitation. In contrast to lodgepole pine systems, area with mortality was linked to precipitation variability even at high beetle populations. Projections from climate models indicate future climate conditions will likely provide favorable conditions for beetle outbreaks within nearly all current whitebark pine habitat in the GYE by the middle of this century. Therefore, when surviving and regenerating trees reach ages suitable for beetle attack, there is strong potential for continued whitebark pine mortality due to mountain pine beetle.

Tree mortality from an exceptional drought spanning mesic to semiarid ecoregions

Abstract: Significant areas of the southern USA periodically experience intense drought that can lead to episodic tree mortality events. Because drought tolerance varies among species and size of trees, such events can alter the structure and function of terrestrial ecosystem in ways that are difficult to detect with local data sets or solely with remote-sensing platforms. We investigated a widespread tree mortality event that resulted from the worst 1-year drought on record for the state of Texas, USA. The drought affected ecoregions spanning mesic to semiarid climate zones and provided a unique opportunity to test hypotheses related to how trees of varying genus and size were affected. The study was based on an extensive set of 599 distributed plots, each 0.16 ha, surveyed in the summer following the drought. In each plot, dead trees larger than 12.7 cm in diameter were counted, sized, and identified to the genus level. Estimates of total mortality were obtained for each of 10 regions using a combination of design-based estimators and calibrated remote sensing using MODIS 1-yr change in normalized difference vegetation index products developed by the U.S. Forest Service. As compared with most of the publicized extreme die-off events, this study documents relatively low rates of mortality occurring over a very large area. However, statewide, regional tree mortality was massive, with an estimated 6.2% of the live trees perishing, nearly nine times greater than normal annual mortality. Dead tree diameters averaged larger than the live trees for most ecoregions, and this trend was most pronounced in the wetter climate zones, suggesting a potential re-ordering of species dominance and downward trend in tree size that was specific to climatic regions. The net effect on carbon storage was estimated to be a redistribution of 24-30 Tg C from the live tree to dead tree carbon pool. The dead tree survey documented drought mortality in more than 29 genera across all regions, and surprisingly, drought resistant and sensitive species fared similarly in some regions. Both angiosperms and gymnosperms were affected. These results highlight that drought-driven mortality alters forest structure differently across climatic regions and genera.

Are cattle surrogate wildlife? Savanna plant community composition explained by total herbivory more than herbivore type

Abstract: The widespread replacement of wild ungulate herbivores by domestic livestock in African savannas is composed of two interrelated phenomena: (1) loss or reduction in numbers of individual wildlife species or guilds and (2) addition of livestock to the system. Each can have important implications for plant community dynamics. Yet very few studies have experimentally addressed the individual, combined, and potentially interactive effects of wild vs. domestic herbivore species on herbaceous plant communities within a single system. Additionally, there is little information about whether, and in which contexts, livestock might functionally replace native herbivore wildlife or, alternatively, have fundamentally different effects on plant species composition. The Kenya Long-term Exclosure Experiment, which has been running since 1995, is composed of six treatment combinations of mega-herbivores, meso-herbivore ungulate wildlife, and cattle. We sampled herbaceous vegetation 25 times between 1999 and 2013. We used partial redundancy analysis and linear mixed models to assess effects of herbivore treatments on overall plant community composition and key plant species. Plant communities in the six different herbivore treatments shifted directionally over time and diverged from each other substantially by 2013. Plant community composition was strongly related (R2 = 0.92) to residual plant biomass, a measure of herbivore utilization. Addition of any single herbivore type (cattle, wildlife, or mega-herbivores) caused a shift in plant community composition that was proportional to its removal of plant biomass. These results suggest that overall herbivory pressure, rather than herbivore type or complex interactions among different herbivore types, was the main driver of changes in plant community composition. Individual plant species, however, did respond most strongly to either wild ungulates or cattle. Although these results suggest considerable functional similarity between a suite of native wild herbivores (which included grazers, browsers, and mixed feeders) and cattle (mostly grazers) with respect to understory plant community composition, responses of individual plant species demonstrate that at the plant-population-level impacts of a single livestock species are not functionally identical to those of a diverse group of native herbivores.

Estimating population density and connectivity of American mink using spatial capture–recapture

Abstract: Estimating the abundance or density of populations is fundamental to the conservation and management of species, and as landscapes become more fragmented, maintaining landscape connectivity has become one of the most important challenges for biodiversity conservation. Yet these two issues have never been formally integrated together in a model that simultaneously models abundance while accounting for connectivity of a landscape. We demonstrate an application of using capture–recapture to develop a model of animal density using a least-cost path model for individual encounter probability that accounts for non-Euclidean connectivity in a highly structured network. We utilized scat detection dogs (Canis lupus familiaris) as a means of collecting non-invasive genetic samples of American mink (Neovison vison) individuals and used spatial capture–recapture models (SCR) to gain inferences about mink population density and connectivity. Density of mink was not constant across the landscape, but rather increased with increasing distance from city, town, or village centers, and mink activity was associated with water. The SCR model allowed us to estimate the density and spatial distribution of individuals across a 388 km2 area. The model was used to investigate patterns of space usage and to evaluate covariate effects on encounter probabilities, including differences between sexes. This study provides an application of capture–recapture models based on ecological distance, allowing us to directly estimate landscape connectivity. This approach should be widely applicable to provide simultaneous direct estimates of density, space usage, and landscape connectivity for many species.

Quantifying dilution and amplification in a community of hosts for tick-borne pathogens

Abstract: Recent controversy over whether biodiversity reduces disease risk (dilution effect) has focused on the ecology of Lyme disease, a tick-borne zoonosis. A criticism of the dilution effect is that increasing host species richness might amplify disease risk, assuming that total host abundance, and therefore feeding opportunities for ticks, increase with species richness. In contrast, a dilution effect is expected when poor quality hosts for ticks and pathogens (dilution hosts) divert tick blood meals away from competent hosts. Even if host densities are additive, the relationship between host density and tick encounters can be nonlinear if the number of ticks that encounter a host is a saturating function of host density, which occurs if ticks aggregate on the remaining hosts rather than failing to find a host before death. Both dilution and amplification are theoretical possibilities, and assessing which is more prevalent required detailed analyses of empirical systems. We used field data to explore the degree of tick redistribution onto fewer individuals with variation in intraspecific host density and novel data-driven models for tick dynamics to determine how changes in vertebrate community composition influence the density of nymphs infected with the Lyme bacterium. To be conservative, we allowed total host density to increase additively with species richness. Our long-term field studies found that larval and nymphal ticks redistribute onto fewer individuals as host densities decline, that a large proportion of nymphs and adults find hosts, and that mice and chipmunks feed a large proportion of nymphs. White-footed mice, eastern chipmunks, short-tailed shrews, and masked shrews were important amplification hosts that greatly increased the density of infected nymphs. Gray squirrels and Virginia opossums were important dilution hosts. Removing these two species increased the maximum number of larvae attached to amplification hosts by 57%. Raccoons and birds were minor dilution hosts under some conditions. Even under the assumption of additive community assembly, some species are likely to reduce the density of infected nymphs as diversity increases. If the assumption of additivity is relaxed, then species that reduce the density of small mammals through predation or competition might substantially reduce disease risk.

Ecologically sustainable weed management: How do we get from proof-of-concept to adoption?

Abstract: Weed management is a critically important activity on both agricultural and non-agricultural lands, but it is faced with a daunting set of challenges: environmental damage caused by control practices, weed resistance to herbicides, accelerated rates of weed dispersal through global trade, and greater weed impacts due to changes in climate and land use. Broad-scale use of new approaches is needed if weed management is to be successful in the coming era. We examine three approaches likely to prove useful for addressing current and future challenges from weeds: diversifying weed management strategies with multiple complementary tactics, developing crop genotypes for enhanced weed suppression, and tailoring management strategies to better accommodate variability in weed spatial distributions. In all three cases, proof-of-concept has long been demonstrated and considerable scientific innovations have been made, but uptake by farmers and land managers has been extremely limited. Impediments to employing these and other ecologically based approaches include inadequate or inappropriate government policy instruments, a lack of market mechanisms, and a paucity of social infrastructure with which to influence learning, decision-making, and actions by farmers and land managers. We offer examples of how these impediments are being addressed in different parts of the world, but note that there is no clear formula for determining which sets of policies, market mechanisms, and educational activities will be effective in various locations. Implementing new approaches for weed management will require multidisciplinary teams comprised of scientists, engineers, economists, sociologists, educators, farmers, land managers, industry personnel, policy makers, and others willing to focus on weeds within whole farming systems and land management units.

Assessing the umbrella value of a range-wide conservation network for jaguars (Panthera onca).

Abstract: Umbrella species are employed as conservation short-cuts for the design of reserves or reserve networks. However, empirical data on the effectiveness of umbrellas is equivocal, which has prevented more widespread application of this conservation strategy. We perform a novel, large-scale evaluation of umbrella species by assessing the potential umbrella value of a jaguar (Panthera onca) conservation network (consisting of viable populations and corridors) that extends from Mexico to Argentina. Using species richness, habitat quality, and fragmentation indices of ~1500 co-occurring mammal species, we show that jaguar populations and corridors overlap a substantial amount and percentage of high-quality habitat for co-occurring mammals and that the jaguar network performs better than random networks in protecting high-quality, interior habitat. Significantly, the effectiveness of the jaguar network as an umbrella would not have been noticeable had we focused on species richness as our sole metric of umbrella utility. Substantial inter-order variability existed, indicating the need for complementary conservation strategies for certain groups of mammals. We offer several reasons for the positive result we document, including the large spatial scale of our analysis and our focus on multiple metrics of umbrella effectiveness. Taken together, our results demonstrate that a regional, single-species conservation strategy can serve as an effective umbrella for the larger community and should help conserve viable populations and connectivity for a suite of co-occurring mammals. Current and future range-wide planning exercises for other large predators may therefore have important umbrella benefits.

Quantifying “apparent” impact and distinguishing impact from invasiveness in multispecies plant invasions

Abstract: The quantification of invader impacts remains a major hurdle to understanding and managing invasions. Here, we demonstrate a method for quantifying the community-level impact of multiple plant invaders by applying Parker et al.'s (1999) equation (impact = range x local abundance x per capita effect or per unit effect) using data from 620 survey plots from 31 grasslands across west-central Montana, USA. In testing for interactive effects of multiple invaders on native plant abundance (percent cover), we found no evidence for invasional meltdown or synergistic interactions for the 25 exotics tested. While much concern exists regarding impact thresholds, we also found little evidence for nonlinear relationships between invader abundance and impacts. These results suggest that management actions that reduce invader abundance should reduce invader impacts monotonically in this system. Eleven of 25 invaders had significant per unit impacts (negative local-scale relationships between invader and native cover). In decomposing the components of impact, we found that local invader abundance had a significant influence on the likelihood of impact, but range (number of plots occupied) did not. This analysis helped to differentiate measures of invasiveness (local abundance and range) from impact to distinguish high-impact invaders from invaders that exhibit negligible impacts, even when widespread. Distinguishing between high- and low-impact invaders should help refine trait-based prediction of problem species. Despite the unique information derived from evaluation of per unit effects of invaders, invasiveness 'scores based on range and local abundance produced similar rankings to impact scores that incorporated estimates of per unit effects. Hence, information on range and local abundance alone was sufficient to identify problematic plant invaders at the regional scale. In comparing empirical data on invader impacts to the state noxious weed list, we found that the noxious weed list captured 45% of the high impact invaders but missed 55% and assigned the lowest risk category to the highest-impact invader. While such subjective weed lists help to guide invasive species management, empirical data are needed to develop more comprehensive rankings of ecological impacts. Using weed lists to classify invaders for testing invasion theory is not well supported.