Feeding en route: Prey availability and traits influence prey selection by an avian predator on migration.

During animal migration, ephemeral communities of taxa at all trophic levels co-occur over space and time. The interactions between predators and prey along migration corridors are ecologically and evolutionarily significant. However, these interactions remain understudied in terrestrial systems and warrant further investigations using novel approaches. We investigated the predator-prey interactions between a migrating avivorous predator and ephemeral avian prey community in the fall migration season. We tested for associations between avian traits and prey selection and hypothesized that prey traits (i.e. relative size, flocking behaviour, habitat, migration tendency and availability) would influence prey selection by a sexually dimorphic raptor on migration. To document prey consumption, we sampled trace prey DNA from beaks and talons of migrating sharp-shinned hawks Accipiter striatus (n = 588). We determined prey availability in the ephemeral avian community by extracting weekly abundance indices from eBird Status and Trends data. We used discrete choice models to assess prey selection and visualized the frequency of prey in diet and availability on the landscape over the fall migration season. Using eDNA metabarcoding, we detected prey species on 94.1% of the hawks sampled (n = 525/588) comprising 1396 prey species detections from 65 prey species. Prey frequency in diet and eBird relative abundance of prey species were correlated over the migration season for top-selected prey species, suggesting prey availability is an important component of raptor-songbird interactions during fall. Prey size, flocking behaviour and non-breeding habitat association were prey traits that significantly influenced predator choice. We found differences between female and male hawk prey selection, suggesting that sexual size dimorphism has led to distinct foraging strategies on migration. This research integrated field data collected by a volunteer-powered raptor migration monitoring station and public-generated data from eBird to reveal elusive predator-prey dynamics occurring in an ephemeral raptor-songbird community during fall migration. Understanding dynamic raptor-songbird interactions along migration routes remains a relatively unexplored frontier in animal ecology and is necessary for the conservation and management efforts of migratory and resident communities.

Durante la migración animal, las comunidades efímeras de taxones de todos los niveles tróficos coexisten en el espacio y el tiempo. Las interacciones entre depredadores y presas a lo largo de los corredores migratorios son significativas desde el punto de vista ecológica y evolutivo. Sin embargo, estas interacciones siguen siendo poco estudiadas en los sistemas terrestres y justifican más investigaciones utilizando enfoques novedosos. Investigamos las interacciones depredador­presa entre un depredador avívoro migratorio y una comunidad de presas aviares efímeras en la temporada migratoria otoñal. Probamos las asociaciones entre los rasgos de las aves y la selección de presas y planteamos la hipótesis de que los rasgos de las presas (tamaño relativo, comportamiento de bandada, hábitat, tendencia migratoria y disponibilidad) influirían en la selección de presas por parte de una rapaz sexualmente dimórfica durante la migración. Para documentar el consumo de presas, recogimos rastros de ADN de presas de picos y garras de Gavilán Americano Accipiter striatus (n = 588) migratorios. Determinamos la disponibilidad de presas en la comunidad de aves efímeras extrayendo índices de abundancia semanales de los datos de eBird Estado y Tendencias. Utilizamos modelos de elección discreta para evaluar la selección de presas y visualizamos la frecuencia de las presas en la dieta y la disponibilidad en el paisaje durante la temporada migratoria otoñal. Utilizando el metacódigo de barras del ADN ambiental, detectamos especies de presas en el 94,1% de los halcones muestreados (n = 525/588), comprendiendo 1396 detecciones de 65 especies de presas. La frecuencia de presas en la dieta y la abundancia relativa de especies de presas en eBird se correlacionaron a lo largo de la temporada de migración para las principales especies de presas seleccionadas, lo que sugiere que la disponibilidad de presas es un componente importante de las interacciones entre aves rapaces y aves canoras durante el otoño. El tamaño de las presas, el comportamiento de las bandadas y la asociación con el hábitat no reproductivo fueron rasgos de presa que influyeron significativamente en la elección de los depredadores. Encontramos diferencias entre la selección de presas de gavilán hembra y macho, lo que sugiere que el dimorfismo sexual de tamaño ha conducido a distintas estrategias de alimentación durante la migración. Esta investigación integró datos de campo recopilados por una estación de monitoreo de migración de rapaces impulsada por voluntarios y datos generados públicamente por eBird para revelar la esquiva dinámica depredador­presa que ocurre en una comunidad efímera de rapaces y aves canoras durante la migración otoñal. Comprender las interacciones dinámicas entre rapaces y aves canoras a lo largo de las rutas migratorias sigue siendo una frontera relativamente inexplorada en la ecología animal y es necesaria para los esfuerzos de conservación y gestión de las comunidades migratorias y residentes.

Strategic restoration planning for land birds in the Colorado River Delta, Mexico.

Ecological restoration is an essential strategy for mitigating the current biodiversity crisis, yet restoration actions are costly. We used systematic conservation planning principles to design an approach that prioritizes restoration sites for birds and tested it in a riparian forest restoration program in the Colorado River Delta. Restoration goals were to maximize the abundance and diversity of 15 priority birds with a variety of habitat preferences. We built abundance models for priority birds based on the current landscape, and predicted bird distributions and relative abundances under a scenario of complete riparian forest restoration throughout our study area. Then, we used Zonation conservation planning software to rank this restored landscape based on core areas for all priority birds. The locations with the highest ranks represented the highest priorities for restoration and were located throughout the river reach. We optimized how much of the available landscape to restore by simulating restoration of the top 10-90% of ranked sites in 10% intervals. We found that total diversity was maximized when 40% of the landscape was restored, and mean relative abundance was maximized when 80% of the landscape was restored. The results suggest that complete restoration is not optimal for this community of priority birds and restoration of approximately 60% of the landscape would provide a balance between maximum relative abundance and diversity. Subsequent planning efforts will combine our results with an assessment of restoration costs to provide further decision support for the restoration-siting process. Our approach can be applied to any landscape-scale restoration program to improve the return on investment of limited economic resources for restoration.

Opposing global change drivers counterbalance trends in breeding North American monarch butterflies.

Many insects are in clear decline, with monarch butterflies (Danaus plexippus) drawing particular attention as a flagship species. It is well documented that, among migratory populations, numbers of overwintering monarchs have been falling across several decades, but trends among breeding monarchs are less clear. Here, we compile >135,000 monarch observations between 1993 and 2018 from the North American Butterfly Association's annual butterfly count to examine spatiotemporal patterns and potential drivers of adult monarch relative abundance trends across the entire breeding range in eastern and western North America. While the data revealed declines at some sites, particularly the US Northeast and parts of the Midwest, numbers in other areas, notably the US Southeast and Northwest, were unchanged or increasing, yielding a slightly positive overall trend across the species range. Negative impacts of agricultural glyphosate use appeared to be counterbalanced by positive effects of annual temperature, particularly in the US Midwest. Overall, our results suggest that population growth in summer is compensating for losses during the winter and that changing environmental variables have offsetting effects on mortality and/or reproduction. We suggest that density-dependent reproductive compensation when lower numbers arrive each spring is currently able to maintain relatively stable breeding monarch numbers. However, we caution against complacency since accelerating climate change may bring growing threats. In addition, increases of summer monarchs in some regions, especially in California and in the south, may reflect replacement of migratory with resident populations. Nonetheless, it is perhaps reassuring that ubiquitous downward trends in summer monarch abundance are not evident.

Integrating data types to estimate spatial patterns of avian migration across the Western Hemisphere.

For many avian species, spatial migration patterns remain largely undescribed, especially across hemispheric extents. Recent advancements in tracking technologies and high-resolution species distribution models (i.e., eBird Status and Trends products) provide new insights into migratory bird movements and offer a promising opportunity for integrating independent data sources to describe avian migration. Here, we present a three-stage modeling framework for estimating spatial patterns of avian migration. First, we integrate tracking and band re-encounter data to quantify migratory connectivity, defined as the relative proportions of individuals migrating between breeding and nonbreeding regions. Next, we use estimated connectivity proportions along with eBird occurrence probabilities to produce probabilistic least-cost path (LCP) indices. In a final step, we use generalized additive mixed models (GAMMs) both to evaluate the ability of LCP indices to accurately predict (i.e., as a covariate) observed locations derived from tracking and band re-encounter data sets versus pseudo-absence locations during migratory periods and to create a fully integrated (i.e., eBird occurrence, LCP, and tracking/band re-encounter data) spatial prediction index for mapping species-specific seasonal migrations. To illustrate this approach, we apply this framework to describe seasonal migrations of 12 bird species across the Western Hemisphere during pre- and postbreeding migratory periods (i.e., spring and fall, respectively). We found that including LCP indices with eBird occurrence in GAMMs generally improved the ability to accurately predict observed migratory locations compared to models with eBird occurrence alone. Using three performance metrics, the eBird + LCP model demonstrated equivalent or superior fit relative to the eBird-only model for 22 of 24 species-season GAMMs. In particular, the integrated index filled in spatial gaps for species with over-water movements and those that migrated over land where there were few eBird sightings and, thus, low predictive ability of eBird occurrence probabilities (e.g., Amazonian rainforest in South America). This methodology of combining individual-based seasonal movement data with temporally dynamic species distribution models provides a comprehensive approach to integrating multiple data types to describe broad-scale spatial patterns of animal movement. Further development and customization of this approach will continue to advance knowledge about the full annual cycle and conservation of migratory birds.

Unraveling a century of global change impacts on winter bird distributions in the eastern United States.

One of the most pressing questions in ecology and conservation centers on disentangling the relative impacts of concurrent global change drivers, climate and land-use/land-cover (LULC), on biodiversity. Yet studies that evaluate the effects of both drivers on species' winter distributions remain scarce, hampering our ability to develop full-annual-cycle conservation strategies. Additionally, understanding how groups of species differentially respond to climate versus LULC change is vital for efforts to enhance bird community resilience to future environmental change. We analyzed long-term changes in winter occurrence of 89 species across nine bird groups over a 90-year period within the eastern United States using Audubon Christmas Bird Count (CBC) data. We estimated variation in occurrence probability of each group as a function of spatial and temporal variation in winter climate (minimum temperature, cumulative precipitation) and LULC (proportion of group-specific and anthropogenic habitats within CBC circle). We reveal that spatial variation in bird occurrence probability was consistently explained by climate across all nine species groups. Conversely, LULC change explained more than twice the temporal variation (i.e., decadal changes) in bird occurrence probability than climate change on average across groups. This pattern was largely driven by habitat-constrained species (e.g., grassland birds, waterbirds), whereas decadal changes in occurrence probabilities of habitat-unconstrained species (e.g., forest passerines, mixed habitat birds) were equally explained by both climate and LULC changes over the last century. We conclude that climate has generally governed the winter occurrence of avifauna in space and time, while LULC change has played a pivotal role in driving distributional dynamics of species with limited and declining habitat availability. Effective land management will be critical for improving species' resilience to climate change, especially during a season of relative resource scarcity and critical energetic trade-offs.

Effects of stewardship on protected area effectiveness for coastal birds.

Evaluation of protected area effectiveness is critical for conservation of biodiversity. Protected areas that prioritize biodiversity conservation are, optimally, located and managed in ways that support relatively large and stable or increasing wildlife populations. Yet evaluating conservation efficacy remains a challenging endeavor. We used an extensive community science data set, eBird, to evaluate the efficacy of protected areas for birds across the Gulf of Mexico and Atlantic coasts of the United States. We modeled trends (2007-2018) for 12 vulnerable waterbirds that use coastal areas during breeding or wintering. We compared two types of protected areas-sites where conservation organizations implemented active stewardship or management or both to reduce human disturbance (hereafter stewardship sites) and local, state, federal, and private protected areas managed to maintain natural land cover (hereafter protected areas)-as well as unprotected areas. We evaluated differences in trends between stewardship, protected, and unprotected areas across the Gulf and Atlantic coasts as a whole. Similar to a background sample, stewardship was known to occur at stewardship sites, but unknown at protected and unprotected areas. Four of 12 target species-Black Skimmer (Rynchops niger), Brown Pelican (Pelecanus occidentalis), Least Tern (Sternula antillarum), and Piping Plover (Charadrius melodus)-had more positive trends (two to 34 times greater) at stewardship sites than protected areas. Furthermore, five target species showed more positive trends at sites with stewardship programs than unprotected sites during at least one season, whereas seven species showed more positive trends at protected than unprotected areas. No species had more negative trends at stewardship sites than unprotected areas, and two species had more negative trends at protected than unprotected areas. Anthropogenic disturbance is a serious threat to coastal birds, and our findings demonstrate that stewardship to reduce its negative impacts helps ensure conservation of vulnerable waterbirds.

La evaluación de la efectividad de las áreas protegidas es de suma importancia para la conservación de la biodiversidad. Las áreas protegidas que priorizan la conservación de la biodiversidad están, de manera óptima, ubicadas y manejadas de maneras que permiten el mantenimiento de poblaciones silvestres relativamente grandes y estables o en incremento. Aun así, la evaluación de la eficacia de la conservación todavía es un esfuerzo desafiante. Usamos un conjunto extensivo de datos de ciencia comunitaria, eBird, para evaluar la eficacia de las áreas protegidas a lo largo de las costas del Golfo de México y del Atlántico en los Estados Unidos. Modelamos las tendencias poblacionales (2007-2018) para doce aves acuáticas vulnerables que usan las áreas costeras durante la temporada de reproducción o de hibernación. Comparamos dos tipos de áreas protegidas - sitios en donde las organizaciones de conservación implementaron una gestión o manejo activo o ambos para reducir la perturbación humana (de ahora en adelante sitios de gestión) y las áreas protegidas locales, estatales, federales y privadas manejadas para mantener la cobertura natural del suelo (de ahora en adelante áreas protegidas) - así como las áreas desprotegidas. Evaluamos las diferencias en las tendencias entre las áreas de gestión, las protegidas y las desprotegidas a lo largo de las cosas del Golfo y del Atlántico como un todo. Similar a una muestra de fondo, se supo que la gestión ocurría en los sitios de gestión, pero no se sabía si ocurría en las áreas protegidas y desprotegidas. Cuatro de las doce especies analizadas - Rhynchops niger, Pelecanus occidentalis, Sternula antillarum y Charadrius melodus - tuvieron tendencias más positivas (2-34 veces mayor) en los sitios de gestión que en las áreas protegidas. Además, cinco especies mostraron más tendencias positivas en los sitios con programas de gestión que en los sitios desprotegidos al menos durante una temporada, mientras que siete especies mostraron más tendencias positivas en sitios protegidos que en las áreas desprotegidas. protegidas que en las desprotegidas. Ninguna especie tuvo más tendencias negativas en los sitios de gestión que en las áreas desprotegidas y dos especies tuvieron más tendencias negativas en las áreas protegidas que en las desprotegidas. La perturbación antropogénica es una amenaza seria para las aves costeras y nuestros hallazgos demuestran que la gestión para reducir sus impactos negativos ayuda a asegurar la conservación de aves acuáticas vulnerables.

Crop pests and predators exhibit inconsistent responses to surrounding landscape composition.

The idea that noncrop habitat enhances pest control and represents a win-win opportunity to conserve biodiversity and bolster yields has emerged as an agroecological paradigm. However, while noncrop habitat in landscapes surrounding farms sometimes benefits pest predators, natural enemy responses remain heterogeneous across studies and effects on pests are inconclusive. The observed heterogeneity in species responses to noncrop habitat may be biological in origin or could result from variation in how habitat and biocontrol are measured. Here, we use a pest-control database encompassing 132 studies and 6,759 sites worldwide to model natural enemy and pest abundances, predation rates, and crop damage as a function of landscape composition. Our results showed that although landscape composition explained significant variation within studies, pest and enemy abundances, predation rates, crop damage, and yields each exhibited different responses across studies, sometimes increasing and sometimes decreasing in landscapes with more noncrop habitat but overall showing no consistent trend. Thus, models that used landscape-composition variables to predict pest-control dynamics demonstrated little potential to explain variation across studies, though prediction did improve when comparing studies with similar crop and landscape features. Overall, our work shows that surrounding noncrop habitat does not consistently improve pest management, meaning habitat conservation may bolster production in some systems and depress yields in others. Future efforts to develop tools that inform farmers when habitat conservation truly represents a win-win would benefit from increased understanding of how landscape effects are modulated by local farm management and the biology of pests and their enemies.

Effects of Elevated Atmospheric Carbon Dioxide and Tropospheric Ozone on Phytochemical Composition of Trembling Aspen ( Populus tremuloides ) and Paper Birch ( Betula papyrifera ).

Anthropogenic activities are altering levels of atmospheric carbon dioxide (CO2) and tropospheric ozone (O3). These changes can alter phytochemistry, and in turn, influence ecosystem processes. We assessed the individual and combined effects of elevated CO2 and O3 on the phytochemical composition of two tree species common to early successional, northern temperate forests. Trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) were grown at the Aspen FACE (Free-Air Carbon dioxide and ozone Enrichment) facility under four combinations of ambient and elevated CO2 and O3. We measured, over three years (2006-08), the effects of CO2 and O3 on a suite of foliar traits known to influence forest functioning. Elevated CO2 had minimal effect on foliar nitrogen and carbohydrate levels in either tree species, and increased synthesis of condensed tannins and fiber in aspen, but not birch. Elevated O3 decreased nitrogen levels in both tree species and increased production of sugar, condensed tannins, fiber, and lignin in aspen, but not birch. The magnitude of responses to elevated CO2 and O3 varied seasonally for both tree species. When co-occurring, CO2 offset most of the changes in foliar chemistry expressed under elevated O3 alone. Our results suggest that levels of CO2 and O3 predicted for the mid-twenty-first century will alter the foliar chemistry of northern temperate forests with likely consequences for forest community and ecosystem dynamics.

From pest data to abundance-based risk maps combining eco-physiological knowledge, weather, and habitat variability.

Noxious species, i.e., crop pest or invasive alien species, are major threats to both natural and managed ecosystems. Invasive pests are of special importance, and knowledge about their distribution and abundance is fundamental to minimize economic losses and prioritize management activities. Occurrence models are a common tool used to identify suitable zones and map priority areas (i.e., risk maps) for noxious species management, although they provide a simplified description of species dynamics (i.e., no indication on species density). An alternative is to use abundance models, but translating abundance data into risk maps is often challenging. Here, we describe a general framework for generating abundance-based risk maps using multi-year pest data. We used an extensive data set of 3968 records collected between 2003 and 2013 in Wisconsin during annual surveys of soybean aphid (SBA), an exotic invasive pest in this region. By using an integrative approach, we modelled SBA responses to weather, seasonal, and habitat variability using generalized additive models (GAMs). Our models showed good to excellent performance in predicting SBA occurrence and abundance (TSS = 0.70, AUC = 0.92; R2  = 0.63). We found that temperature, precipitation, and growing degree days were the main drivers of SBA trends. In addition, a significant positive relationship between SBA abundance and the availability of overwintering habitats was observed. Our models showed aphid populations were also sensitive to thresholds associated with high and low temperatures, likely related to physiological tolerances of the insects. Finally, the resulting aphid predictions were integrated using a spatial prioritization algorithm ("Zonation") to produce an abundance-based risk map for the state of Wisconsin that emphasized the spatiotemporal consistency and magnitude of past infestation patterns. This abundance-based risk map can provide information on potential foci of pest outbreaks where scouting efforts and prophylactic measures should be concentrated. The approach we took is general, relatively simple, and can be applied to other species, habitats and geographical areas for which species abundance data and biotic and abiotic data are available.

A Landscape View of Agricultural Insecticide Use across the Conterminous US from 1997 through 2012.

Simplification of agricultural landscapes is expected to have positive effects on many crop pests and negative effects on their natural enemies, potentially leading to increased pest pressure, decreased crop yield, and increased insecticide use. While many intermediate links in this causal chain have empirical support, there is mixed evidence for ultimate relationships between landscape simplification, crop yield, and insecticide use, especially at large spatial and temporal scales. We explored relationships between landscape simplification (proportion of a county in harvested cropland) and insecticide use (proportion of harvested cropland treated with insecticides), using county-level data from the US Census of Agriculture and a variety of standard and spatiotemporal regression techniques. The best model indicated that insecticide use across the US has increased between 1997 and 2012, was strongly dependent on the crops grown in a county, increased with average farm income and size, and increased with annual growing degree days. After accounting for those variables, and other unidentified spatial and temporal structure in the data, there remained a statistically significant, moderate, positive relationship between insecticide use and landscape simplification. These results lend general support to the causal chain outlined above, and to the notion that a landscape perspective is useful for managing ecosystem services that are provided by mobile organisms and valuable to agriculture.

Explicit modeling of abiotic and landscape factors reveals precipitation and forests associated with aphid abundance.

Increases in natural or noncrop habitat surrounding agricultural fields have been shown to be correlated with declines in insect crop pests. However, these patterns are highly variable across studies suggesting other important factors, such as abiotic drivers, which are rarely included in landscape models, may also contribute to variability in insect population abundance. The objective of this study was to explicitly account for the contribution of temperature and precipitation, in addition to landscape composition, on the abundance of a widespread insect crop pest, the soybean aphid (Aphis glycines Matsumura), in Wisconsin soybean fields. We hypothesized that higher soybean aphid abundance would be associated with higher heat accumulation (e.g., growing degree days) and increasing noncrop habitat in the surrounding landscape, due to the presence of the overwintering primary hosts of soybean aphid. To evaluate these hypotheses, we used an ecoinformatics approach that relied on a large dataset collected across Wisconsin over a 9-year period (2003-2011), for an average of 235 sites per year (n = 2,110 fields total). We determined surrounding landscape composition (1.5-km radius) using publicly available satellite-derived land cover imagery and interpolated daily temperature and precipitation information from the National Weather Service COOP weather station network. We constructed linear mixed models for soybean aphid abundance based on abiotic and landscape explanatory variables and applied model averaging for prediction using an information theoretic framework. Over this broad spatial and temporal extent in Wisconsin, we found that variation in growing season precipitation was positively related to soybean aphid abundance, while higher precipitation during the nongrowing season had a negative effect on aphid populations. Additionally, we found that aphid populations were higher in areas with proportionally more forest but were lower in areas where minor crops, such as small grains, were more prevalent. Thus, our findings support our hypothesis that including abiotic drivers increases our understanding of crop pest abundance and distribution. Moreover, by explicitly modeling abiotic factors, we may be able to explore how variable climate in tandem with land cover patterns may affect current and future insect populations, with potentially critical implications for crop yields and agricultural food webs.

Alternative scenarios of bioenergy crop production in an agricultural landscape and implications for bird communities.

Increased demand and government mandates for bioenergy crops in the United States could require a large allocation of agricultural land to bioenergy feedstock production and substantially alter current landscape patterns. Incorporating bioenergy landscape design into land-use decision making could help maximize benefits and minimize trade-offs among alternative land uses. We developed spatially explicit landscape scenarios of increased bioenergy crop production in an 80-km radius agricultural landscape centered on a potential biomass-processing energy facility and evaluated the consequences of each scenario for bird communities. Our scenarios included conversion of existing annual row crops to perennial bioenergy grasslands and conversion of existing grasslands to annual bioenergy row crops. The scenarios explored combinations of four biomass crop types (three potential grassland crops along a gradient of plant diversity and one annual row crop [corn]), three land conversion percentages to bioenergy crops (10%, 20%, or 30% of row crops or grasslands), and three spatial configurations of biomass crop fields (random, clustered near similar field types, or centered on the processing plant), yielding 36 scenarios. For each scenario, we predicted the impact on four bird community metrics: species richness, total bird density, species of greatest conservation need (SGCN) density, and SGCN hotspots (SGCN birds/ha ≥ 2). Bird community metrics consistently increased with conversion of row crops to bioenergy grasslands and consistently decreased with conversion of grasslands to bioenergy row crops. Spatial arrangement of bioenergy fields had strong effects on the bird community and in some cases was more influential than the amount converted to bioenergy crops. Clustering grasslands had a stronger positive influence on the bird community than locating grasslands near the central plant or at random. Expansion of bioenergy grasslands onto marginal agricultural lands will likely benefit grassland bird populations, and bioenergy landscapes could be designed to maximize biodiversity benefits while meeting targets for biomass production.

Evidence for Compensatory Photosynthetic and Yield Response of Soybeans to Aphid Herbivory.

The soybean aphid, Aphis glycines Matsumura, an exotic species in North America that has been detected in 21 U.S. states and Canada, is a major pest for soybean that can reduce maximum photosynthetic capacity and yields. Our existing knowledge is based on relatively few studies that do not span a wide variety of environmental conditions, and often focus on relatively high and damaging population pressure. We examined the effects of varied populations and duration of soybean aphids on soybean photosynthetic rates and yield in two experiments. In a 2011 field study, we found that plants with low cumulative aphid days (CAD, less than 2,300) had higher yields than plants not experiencing significant aphid pressure, suggesting a compensatory growth response to low aphid pressure. This response did not hold at higher CAD, and yields declined. In a 2013 controlled-environment greenhouse study, soybean plants were well-watered and fertilized with nitrogen (N), and aphid populations were manipulated to reach moderate to high levels (8,000-50,000 CAD). Plants tolerated these population levels when aphids were introduced during the vegetative or reproductive phenological stages of the plant, showing no significant reduction in yield. Leaf N concentration and CAD were positively and significantly correlated with increasing ambient photosynthetic rates. Our findings suggest that, given the right environmental conditions, modern soybean plants can withstand higher aphid pressure than previously assumed. Moreover, soybean plants also responded positively through a compensatory photosynthetic effect to moderate population pressure, contributing to stable or increased yield.

Trophic cascades in agricultural landscapes: indirect effects of landscape composition on crop yield.

The strength and prevalence of trophic cascades, defined as positive, indirect effects of natural enemies (predatory and parasitic arthropods) on plants, is highly variable in agroecosystems. This variation may in part be due to the spatial or landscape context in which hese trophic cascades occur. In 2011 and 2012, we conducted a natural enemy exclusion experiment in soybean fields along a gradient of landscape composition across southern Wisconsin and Michigan, USA. We used structural equation modeling to ask (1) whether natural enemies influence biocontrol of soybean aphids (SBA) and soybean yield and (2) whether landscape effects on natural enemies influence the strength of the trophic cascades. We found that natural enemies (NE) suppressed aphid populations in both years of our study, and, in 2011, the yield of soybean plants exposed to natural enemies was 37% higher than the yield of plants with aphid populations protected from natural enemies. The strength of the :rophic cascade was also influenced by landscape context. We found that landscapes with a higher proportion of soybean and higher diversity habitats resulted in more NE, fewer aphids, and, in some cases, a trend toward greater soybean yield. These results indicate that landscape context is important for understanding spatial variability in biocontrol and yield, but other factors, such as environmental variability and compensatory growth, might overwhelm the beneficial effects of biocontrol on crop yield.

Modeling pollinator community response to contrasting bioenergy scenarios.

In the United States, policy initiatives aimed at increasing sources of renewable energy are advancing bioenergy production, especially in the Midwest region, where agricultural landscapes dominate. While policy directives are focused on renewable fuel production, biodiversity and ecosystem services will be impacted by the land-use changes required to meet production targets. Using data from field observations, we developed empirical models for predicting abundance, diversity, and community composition of flower-visiting bees based on land cover. We used these models to explore how bees might respond under two contrasting bioenergy scenarios: annual bioenergy crop production and perennial grassland bioenergy production. In the two scenarios, 600,000 ha of marginal annual crop land or marginal grassland were converted to perennial grassland or annual row crop bioenergy production, respectively. Model projections indicate that expansion of annual bioenergy crop production at this scale will reduce bee abundance by 0 to 71%, and bee diversity by 0 to 28%, depending on location. In contrast, converting annual crops on marginal soil to perennial grasslands could increase bee abundance from 0 to 600% and increase bee diversity between 0 and 53%. Our analysis of bee community composition suggested a similar pattern, with bee communities becoming less diverse under annual bioenergy crop production, whereas bee composition transitioned towards a more diverse community dominated by wild bees under perennial bioenergy crop production. Models, like those employed here, suggest that bioenergy policies have important consequences for pollinator conservation.

Herbivore-mediated material fluxes in a northern deciduous forest under elevated carbon dioxide and ozone concentrations.

Anthropogenic changes in atmospheric carbon dioxide (CO2 ) and ozone (O3 ) are known to alter tree physiology and growth, but the cascading effects on herbivore communities and herbivore-mediated nutrient cycling are poorly understood. We sampled herbivore frass, herbivore-mediated greenfall, and leaf-litter deposition in temperate forest stands under elevated CO2 (c. 560 ppm) and O3 (c. 1.5× ambient), analyzed substrate chemical composition, and compared the quality and quantity of fluxes under multiple atmospheric treatments. Leaf-chewing herbivores fluxed 6.2 g m(-2)  yr(-1) of frass and greenfall from the canopy to the forest floor, with a carbon : nitrogen (C : N) ratio 32% lower than that of leaf litter. Herbivore fluxes of dry matter, C, condensed tannins, and N increased under elevated CO2 (35, 32, 63 and 39%, respectively), while fluxes of N decreased (18%) under elevated O3 . Herbivore-mediated dry matter inputs scaled across atmospheric treatments as a constant proportion of leaf-litter inputs. Increased fluxes under elevated CO2 were consistent with increased herbivore consumption and abundance, and with increased plant growth and soil respiration, previously reported for this experimental site. Results suggest that insect herbivory will reinforce other factors, such as photosynthetic rate and fine-root production, impacting C sequestration by forests in future environments.

Perennial grasslands enhance biodiversity and multiple ecosystem services in bioenergy landscapes.

Agriculture is being challenged to provide food, and increasingly fuel, for an expanding global population. Producing bioenergy crops on marginal lands--farmland suboptimal for food crops--could help meet energy goals while minimizing competition with food production. However, the ecological costs and benefits of growing bioenergy feedstocks--primarily annual grain crops--on marginal lands have been questioned. Here we show that perennial bioenergy crops provide an alternative to annual grains that increases biodiversity of multiple taxa and sustain a variety of ecosystem functions, promoting the creation of multifunctional agricultural landscapes. We found that switchgrass and prairie plantings harbored significantly greater plant, methanotrophic bacteria, arthropod, and bird diversity than maize. Although biomass production was greater in maize, all other ecosystem services, including methane consumption, pest suppression, pollination, and conservation of grassland birds, were higher in perennial grasslands. Moreover, we found that the linkage between biodiversity and ecosystem services is dependent not only on the choice of bioenergy crop but also on its location relative to other habitats, with local landscape context as important as crop choice in determining provision of some services. Our study suggests that bioenergy policy that supports coordinated land use can diversify agricultural landscapes and sustain multiple critical ecosystem services.

Ecosystem-service tradeoffs associated with switching from annual to perennial energy crops in riparian zones of the US Midwest.

Integration of energy crops into agricultural landscapes could promote sustainability if they are placed in ways that foster multiple ecosystem services and mitigate ecosystem disservices from existing crops. We conducted a modeling study to investigate how replacing annual energy crops with perennial energy crops along Wisconsin waterways could affect a variety of provisioning and regulating ecosystem services. We found that a switch from continuous corn production to perennial-grass production decreased annual income provisioning by 75%, although it increased annual energy provisioning by 33%, decreased annual phosphorous loading to surface water by 29%, increased below-ground carbon sequestration by 30%, decreased annual nitrous oxide emissions by 84%, increased an index of pollinator abundance by an average of 11%, and increased an index of biocontrol potential by an average of 6%. We expressed the tradeoffs between income provisioning and other ecosystem services as benefit-cost ratios. Benefit-cost ratios averaged 12.06 GJ of additional net energy, 0.84 kg of avoided phosphorus pollution, 18.97 Mg of sequestered carbon, and 1.99 kg of avoided nitrous oxide emissions for every $1,000 reduction in income. These ratios varied spatially, from 2- to 70-fold depending on the ecosystem service. Benefit-cost ratios for different ecosystem services were generally correlated within watersheds, suggesting the presence of hotspots--watersheds where increases in multiple ecosystem services would come at lower-than-average opportunity costs. When assessing the monetary value of ecosystem services relative to existing conservation programs and environmental markets, the overall value of enhanced services associated with adoption of perennial energy crops was far lower than the opportunity cost. However, when we monitized services using estimates for the social costs of pollution, the value of enhanced services far exceeded the opportunity cost. This disparity between recoverable costs and social value represents a fundamental challenge to expansion of perennial energy crops and sustainable agricultural landscapes.

Pest-suppression potential of midwestern landscapes under contrasting bioenergy scenarios.

Biomass crops grown on marginal soils are expected to fuel an emerging bioenergy industry in the United States. Bioenergy crop choice and position in the landscape could have important impacts on a range of ecosystem services, including natural pest-suppression (biocontrol services) provided by predatory arthropods. In this study we use predation rates of three sentinel crop pests to develop a biocontrol index (BCI) summarizing pest-suppression potential in corn and perennial grass-based bioenergy crops in southern Wisconsin, lower Michigan, and northern Illinois. We show that BCI is higher in perennial grasslands than in corn, and increases with the amount of perennial grassland in the surrounding landscape. We develop an empirical model for predicting BCI from information on energy crop and landscape characteristics, and use the model in a qualitative assessment of changes in biocontrol services for annual croplands on prime agricultural soils under two contrasting bioenergy scenarios. Our analysis suggests that the expansion of annual energy crops onto 1.2 million ha of existing perennial grasslands on marginal soils could reduce BCI between -10 and -64% for nearly half of the annual cropland in the region. In contrast, replacement of the 1.1 million ha of existing annual crops on marginal land with perennial energy crops could increase BCI by 13 to 205% on over half of the annual cropland in the region. Through comparisons with other independent studies, we find that our biocontrol index is negatively related to insecticide use across the Midwest, suggesting that strategically positioned, perennial bioenergy crops could reduce insect damage and insecticide use on neighboring food and forage crops. We suggest that properly validated environmental indices can be used in decision support systems to facilitate integrated assessments of the environmental and economic impacts of different bioenergy policies.

Energetics of thermoregulation by an industrious endotherm.

OBJECTIVES: Thermoregulation by modern industrial humans is unique among endothermic animals, in that it is largely accomplished by controlling the temperature of our external environment. The objective of this study was to view the relationship between thermoregulatory energy use and environmental temperature in modern humans from the perspective of comparative physiology. METHODS: Monthly residential energy use estimates from the US Energy Information Administration were divided by the annual number of American households from the US Census Bureau, giving average monthly energy consumption per American household for the years 2006 through 2010. Monthly energy consumption was then plotted against average monthly temperature across the United States from the National Climatic Data Center. RESULTS: The resulting graph bore a striking resemblance to a classic Scholander-Irving curve, exhibiting clear upper (22°C) and lower (15°C) critical temperatures, and an increase in energy use as temperatures extend above (90 W °C(-1) increase) or below (244 W °C(-1) decrease) those critical temperatures. Allometric equations from comparative physiology indicate that the energetic costs of our current thermoregulatory habits are ∼30 to 50 times those predicted for an endotherm of our size. CONCLUSIONS: Modern humans have redefined what it means to be a homeothermic endotherm, using large quantities of extrametabolic energy to regulate the temperature of our surroundings. Despite this sophistication, the signal of our individual physiology is readily discernible in national data on energy consumption.