Capacity and establishment rules govern the number of nonnative species in communities of ground-dwelling invertebrates.

Nonnative species are a key agent of global change. However, nonnative invertebrates remain understudied at the community scales where they are most likely to drive local extirpations. We use the North American NEON pitfall trapping network to document the number of nonnative species from 51 invertebrate communities, testing four classes of drivers. We sequenced samples using the eDNA from the sample's storage ethanol. We used AICc informed regression to evaluate how native species richness, productivity, habitat, temperature, and human population density and vehicular traffic account for continent-wide variation in the number of nonnative species in a local community. The percentage of nonnatives varied 3-fold among habitat types and over 10-fold (0%-14%) overall. We found evidence for two types of constraints on nonnative diversity. Consistent with Capacity rules (i.e., how the number of niches and individuals reflect the number of species an ecosystem can support) nonnatives increased with existing native species richness and ecosystem productivity. Consistent with Establishment Rules (i.e., how the dispersal rate of nonnative propagules and the number of open sites limits nonnative species richness) nonnatives increased with automobile traffic-a measure of human-generated propagule pressure-and were twice as common in pastures than native grasslands. After accounting for drivers associated with a community's ability to support native species (native species richness and productivity), nonnatives are more common in communities that are regularly seasonally disturbed (pastures and, potentially deciduous forests) and those experiencing more vehicular traffic. These baseline values across the US North America will allow NEON's monitoring mission to document how anthropogenic change-from disturbance to propagule transport, from temperature to trends in local extinction-further shape biotic homogenization.

Beyond "greening" and "browning": Trends in grassland ground cover fractions across Eurasia that account for spatial and temporal autocorrelation.

Grassland ecosystems cover up to 40% of the global land area and provide many ecosystem services directly supporting the livelihoods of over 1 billion people. Monitoring long-term changes in grasslands is crucial for food security, biodiversity conservation, achieving Land Degradation Neutrality goals, and modeling the global carbon budget. Although long-term grassland monitoring using remote sensing is extensive, it is typically based on a single vegetation index and does not account for temporal and spatial autocorrelation, which means that some trends are falsely identified while others are missed. Our goal was to analyze trends in grasslands in Eurasia, the largest continuous grassland ecosystems on Earth. To do so, we calculated Cumulative Endmember Fractions (annual sums of monthly ground cover fractions) derived from MODIS 2002-2020 time series, and applied a new statistical approach PARTS that explicitly accounts for temporal and spatial autocorrelation in trends. We examined trends in green vegetation, non-photosynthetic vegetation, and soil ground cover fractions considering their independent change trajectories and relations among fractions over time. We derived temporally uncorrelated pixel-based trend maps and statistically tested whether observed trends could be explained by elevation, land cover, SPEI3, climate, country, and their combinations, all while accounting for spatial autocorrelation. We found no statistical evidence for a decrease in vegetation cover in grasslands in Eurasia. Instead, there was a significant map-level increase in non-photosynthetic vegetation across the region and local increases in green vegetation with a concomitant decrease in soil fraction. Independent environmental variables affected trends significantly, but effects varied by region. Overall, our analyses show in a statistically robust manner that Eurasian grasslands have changed considerably over the past two decades. Our approach enhances remote sensing-based monitoring of trends in grasslands so that underlying processes can be discerned.

Temperature-habitat interactions constrain seasonal activity in a continental array of pitfall traps.

Activity density (AD), the rate at which animals collectively move through their environment, emerges as the product of a taxon's local abundance and its velocity. We analyze drivers of seasonal AD using 47 localities from the National Ecological Observatory Network (NEON) both to better understand variation in ecosystem rates like pollination and seed dispersal as well as the constraints of using AD to monitor invertebrate populations. AD was measured as volume from biweekly pitfall trap arrays (ml trap-1 14 days-1 ). Pooled samples from 2017 to 2018 revealed AD extrema at most temperatures but with a strongly positive overall slope. However, habitat types varied widely in AD's seasonal temperature sensitivity, from negative in wetlands to positive in mixed forest, grassland, and shrub habitats. The temperature of maximum AD varied threefold across the 47 localities; it tracked the threefold geographic variation in maximum growing season temperature with a consistent gap of ca. 3°C across habitats, a novel macroecological result. AD holds potential as an effective proxy for investigating ecosystem rates driven by activity. However, our results suggest that its use for monitoring insect abundance is complicated by the many ways that both abundance and velocity are constrained by a locality's temperature and plant physiognomy.

Activity density at a continental scale: What drives invertebrate biomass moving across the soil surface?

Activity density (AD), the rate that an individual taxon or its biomass moves through the environment, is used both to monitor communities and quantify the potential for ecosystem work. The Abundance Velocity Hypothesis posited that AD increases with aboveground net primary productivity (ANPP) and is a unimodal function of temperature. Here we show that, at continental extents, increasing ANPP may have nonlinear effects on AD: increasing abundance, but decreasing velocity as accumulating vegetation interferes with movement. We use 5 yr of data from the NEON invertebrate pitfall trap arrays including 43 locations and four habitat types for a total of 77 habitat-site combinations to evaluate continental drivers of invertebrate AD. ANPP and temperature accounted for one-third to 92% of variation in AD. As predicted, AD was a unimodal function of temperature in forests and grasslands but increased linearly in open scrublands. ANPP yielded further nonlinear effects, generating unimodal AD curves in wetlands, and bimodal curves in forests. While all four habitats showed no AD trends over 5 yr of sampling, these nonlinearities suggest that trends in AD, often used to infer changes in insect abundance, will vary qualitatively across ecoregions.

How and why plant ionomes vary across North American grasslands and its implications for herbivore abundance.

Plant elemental content can vary up to 1,000-fold across grasslands, with implications for the herbivores the plants feed. We contrast the regulation, in grasses and forbs, of 12 elements essential to plants and animals (henceforth plant-essential), 7 essential to animals but not plants (animal-essential) and 6 with no known metabolic function (nonessential). Four hypotheses accounted for up to two thirds of the variation in grass and forb ionomes across 54 North American grasslands. Consistent with the supply-side hypothesis, the plant-essential ionome of both forbs and grasses tracked soil availability. Grass ionomes were more likely to harvest even nonessential elements like Cd and Sr. Consistent with the grazing hypothesis, cattle-grazed grasslands also accumulated a handful of metals like Cu and Cr. Consistent with the NP-catalysis hypothesis, increases in the macronutrients N and P in grasses were associated with higher densities of cofactors like Zn and Cu. The plant-essential elements of forbs, in contrast, consistently varied as per the nutrient-dilution hypothesis-there was a decrease in elemental parts per million with increasing local carbohydrate production. Combined, these data fit a working hypothesis that grasses maintain lower elemental densities and survive on nutrient-poor patches by opportunistically harvesting soil nutrients. In contrast, nutrient-rich forbs use episodes of high precipitation and temperature to build new carbohydrate biomass, raising leaves higher to compete for light, but diluting the nutrient content in every bite of tissue. Herbivores of forbs may thus be particularly prone to increases in pCO2 via nutrient dilution.

Satellite Observations and Malaria: New Opportunities for Research and Applications.

Satellite remote sensing provides a wealth of information about environmental factors that influence malaria transmission cycles and human populations at risk. Long-term observations facilitate analysis of climate-malaria relationships, and high-resolution data can be used to assess the effects of agriculture, urbanization, deforestation, and water management on malaria. New sources of very-high-resolution satellite imagery and synthetic aperture radar data will increase the precision and frequency of observations. Cloud computing platforms for remote sensing data combined with analysis-ready datasets and high-level data products have made satellite remote sensing more accessible to nonspecialists. Further collaboration between the malaria and remote sensing communities is needed to develop and implement useful geospatial data products that will support global efforts toward malaria control, elimination, and eradication.

Nutrient dilution and climate cycles underlie declines in a dominant insect herbivore.

Evidence for global insect declines mounts, increasing our need to understand underlying mechanisms. We test the nutrient dilution (ND) hypothesis-the decreasing concentration of essential dietary minerals with increasing plant productivity-that particularly targets insect herbivores. Nutrient dilution can result from increased plant biomass due to climate or CO2 enrichment. Additionally, when considering long-term trends driven by climate, one must account for large-scale oscillations including El Niño Southern Oscillation (ENSO), North Atlantic Oscillation (NAO), and Pacific Decadal Oscillation (PDO). We combine long-term datasets of grasshopper abundance, climate, plant biomass, and end-of-season foliar elemental content to examine potential drivers of abundance cycles and trends of this dominant herbivore. Annual grasshopper abundances in 16- and 22-y time series from a Kansas prairie revealed both 5-y cycles and declines of 2.1-2.7%/y. Climate cycle indices of spring ENSO, summer NAO, and winter or spring PDO accounted for 40-54% of the variation in grasshopper abundance, mediated by effects of weather and host plants. Consistent with ND, grass biomass doubled and foliar concentrations of N, P, K, and Na-nutrients which limit grasshopper abundance-declined over the same period. The decline in plant nutrients accounted for 25% of the variation in grasshopper abundance over two decades. Thus a warming, wetter, more CO2-enriched world will likely contribute to declines in insect herbivores by depleting nutrients from their already nutrient-poor diet. Unlike other potential drivers of insect declines-habitat loss, light and chemical pollution-ND may be widespread in remaining natural areas.

Bottom-up when it is not top-down: Predators and plants control biomass of grassland arthropods.

We investigate where bottom-up and top-down control regulates ecological communities as a mechanism linking ecological gradients to the geography of consumer abundance and biomass. We use standardized surveys of 54 North American grasslands to test alternate hypotheses predicting 100-fold shifts in the biomass of four common grassland arthropod taxa-Auchenorrhyncha, sucking herbivores, Acrididae, chewing herbivores, Tettigoniidae, omnivores, and Araneae, predators. Bottom-up models predict that consumer biomass tracks plant quantity (e.g. productivity and standing biomass) and quality (nutrient content) and that ectotherm access to food increases with temperature. Each of the focal trophic groups responded differently to these drivers: the biomass of sucking herbivores and omnivores increased with plant biomass; that of chewing herbivores tracked plant quality; and predator biomass did not depend on plant quality, plant quantity or temperature. The Exploitation Ecosystem Hypothesis is a top-down hypothesis that predicts a shift from resource limitation of herbivores when plant production is low, to predator limitation when plant production is high. In grasslands where spider biomass was low, herbivore biomass increased with plant biomass, whereas bottom-up structuring was not evident when spiders were abundant. Furthermore, neither predator biomass nor trophic position (via stable isotope analysis) increased with plant biomass, suggesting predators themselves are top-down limited. Stable isotope analysis revealed that trophic position of the chewing herbivore and omnivore increased significantly with plant biomass, suggesting these groups increased scavenging and meat consumption in grasslands with higher carbohydrate availability. Taken together, our snapshot sampling documents gradients of food web structure across 54 grasslands, consistent with multiple hypotheses of bottom-up and top-down regulation.

The nutritional geography of ants: Gradients of sodium and sugar limitation across North American grasslands.

Sugar and sodium are essential nutrients to above- and below-ground consumers. Unlike other properties of ecological communities such as abundance and richness, we know relatively little about nutritional geography-the sources and supply rates of nutrients, and how and why they vary across communities and ecosystems. Towards a remedy, we present a suite of hypotheses for how sodium and sugary exudate availability should vary for a common omnivore-the ants-and test them using a survey of 53 North American grasslands. We do so by running transects of salt and sugar baits and inferring the magnitude of environmental supplies as the inverse of their use as exogenous baits. We then use estimates of potential drivers of the availability of salt and sugary exudates-plant and soil nutrients, and bioclimatic variables-to test the best predictors of sodium and salt use by ant communities. Beyond a baseline of ant activity, salt use increased as an inverse of the amount of sodium in an ecosystem's plant tissue, but not its soils. Plant sodium varied by two orders of magnitude in grasslands across 16° latitude. This suggests that plant exudates are an important source of sodium for grassland consumers. The three drivers that best predict exogenous sugar use by ants all point to factors constraining sugar production: net above-ground productivity, how far the community is into that year's growing season (both reflecting the rates of photosynthesis) and, intriguingly, the potassium content of plant tissue, which is likely linked to exudate production via plant turgor. These data suggest that ants and other consumers across a range of grasslands and climate vary significantly in the demand and supply of sugar and salt. This nutritional geography ultimately arises from gradients of climate and biogeochemistry with implications for the geography of plant-consumer interactions.

Species energy and Thermal Performance Theory predict 20-yr changes in ant community abundance and richness.

In an era of rapid climate change, and with it concern over insect declines, we used two theories to predict 20-yr changes in 34 North American ant communities. The ecosystems, from deserts to hardwood forests, were first surveyed in the 1990s. When resurveyed in 2016-2017, they averaged 1°C warmer with 200 g C·m-2 ·yr-1 higher plant productivity. Ant colony abundance changed from -49% to +61%. Consistent with Thermal Performance Theory, colony abundance increased with temperature increases < 1°C, then decreased as a site's mean monthly temperature change increased up to +2.4°C. Consistent with Species Energy Theory, (1) ant abundance tracked changes in a measure of energy availability (net aboveground productivity, g C·m-2 ·yr-1 ) and (2) increases in colony abundance drove increases in local plot- and transect-level species richness but not that of Chao 2, an estimate of the size of the species pool. Even after accounting for these drivers, local species richness was still higher ~20 yr after the original surveys, likely due to the increased activity of ant workers. These results suggest community changes are predictable using theory from geographical ecology, and that warming can first enhance but may ultimately decrease the abundance of this important insect taxon.

A distributed experiment demonstrates widespread sodium limitation in grassland food webs.

Sodium (Na) has a unique role in food webs as a nutrient primarily limiting for plant consumers, but not other trophic levels. Environmental Na levels vary with proximity to coasts, local geomorphology, climate, and with anthropogenic inputs (e.g., road salt). We tested two key predictions across 54 grasslands in North America: Na shortfall commonly limits herbivore abundance, and the magnitude of this limitation varies inversely with environmental Na supplies. We tested them with a distributed pulse experiment and evaluated the relative importance of Na limitation to other classic drivers of climate, macronutrient levels, and plant productivity. Herbivore abundance increased by 45% with Na addition. Moreover, the magnitude of increase on Na addition plots decreased with increasing levels of plant Na, indicating Na satiation at sites with high Na concentrations in plant tissue. Our results demonstrate that invertebrate primary consumers are often Na limited and track local Na availability, with implications for the geography of invertebrate abundance and herbivory.

Evaluation of geoimputation strategies in a large case study.

BACKGROUND: Health data usually has missing or incomplete location information, which impacts the quality of research. Geoimputation methods are used by health professionals to increase the spatial resolution of address information for more accurate analyses. The objective of this study was to evaluate geo-imputation methods with respect to the demographic and spatial characteristics of the data. METHODS: We evaluated four geoimputation methods for increasing spatial resolution of records with known locational information at a coarse level. In order to test and rigorously evaluate two stochastic and two deterministic strategies, we used the Texas Sex Offender registry database with over 50,000 records with known demographic and coordinate information. We reduced the spatial resolution of each record to a census block group and attempted to recover coordinate information using the four strategies. We rigorously evaluated the results in terms of the error distance between the original coordinates and recovered coordinates by studying the results by demographic sub groups and the characteristics of the underlying geography. RESULTS: We observed that in estimating the actual location of a case, the weighted mean method is the most superior for each demographic group followed by the maximum imputation centroid, the random point in matching sub-geographies and the random point in all sub-geographies methods. Higher accuracies were observed for minority populations because minorities tend to cluster in certain neighborhoods, which makes it easier to impute their location. Results are greatly affected by the population density of the underlying geographies. We observed high accuracies in high population density areas, which often exist within smaller census blocks, which makes the search space smaller. Similarly, mapping geoimputation accuracies in a spatially explicit manner reveals that metropolitan areas yield higher accuracy results. CONCLUSIONS: Based on gains in standard error, reduction in mean error and validation results, we conclude that characteristics of the estimated records such as the demographic profile and population density information provide a measure of certainty of geographic imputation.

The Anthropocene Biosphere: Supporting 'Open Interdisciplinarity' through Blogging.

This paper describes a process of 'open' interdisciplinary scholarship. Researchers from across the University of Oklahoma blogged about a recent paper by ecologist Erle Ellis, and met in person to discuss posts. They then hosted Ellis for a seminar on questions that emerged, and for a public panel discussion.

Revealing turning points in ecosystem functioning over the Northern Eurasian agricultural frontier.

The collapse of the Soviet Union in 1991 has been a turning point in the World history that left a unique footprint on the Northern Eurasian ecosystems. Conducting large scale mapping of environmental change and separating between naturogenic and anthropogenic drivers is a difficult endeavor in such highly complex systems. In this research a piece-wise linear regression method was used for breakpoint detection in Rain-Use Efficiency (RUE) time series and a classification of ecosystem response types was produced. Supported by earth observation data, field data, and expert knowledge, this study provides empirical evidence regarding the occurrence of drastic changes in RUE (assessment of the timing, the direction and the significance of these changes) in Northern Eurasian ecosystems between 1982 and 2011. About 36% of the study area (3.4 million km(2) ) showed significant (P < 0.05) trends and/or turning points in RUE during the observation period. A large proportion of detected turning points in RUE occurred around the fall of the Soviet Union in 1991 and in the following years which were attributed to widespread agricultural land abandonment. Our study also showed that recurrent droughts deeply affected vegetation productivity throughout the observation period, with a general worsening of the drought conditions in recent years. Moreover, recent human-induced turning points in ecosystem functioning were detected and attributed to ongoing recultivation and change in irrigation practices in the Volgograd region, and to increased salinization and increased grazing intensity around Lake Balkhash. The ecosystem-state assessment method introduced here proved to be a valuable support that highlighted hotspots of potentially altered ecosystems and allowed for disentangling human from climatic disturbances.

Long-term impacts of land cover changes on stream channel loss.

Land cover change and stream channel loss are two related global environmental changes that are expanding and intensifying. Here, we examine how different types and transitions of land cover change impact stream channel loss across a large urbanizing watershed. We present historical land cover in the 666-km(2) Lake Thunderbird watershed in central Oklahoma (USA) over a 137 year period and coinciding stream channel length changes for the most recent 70 years of this period. Combining these two datasets allowed us to assess the interaction of land cover changes with stream channel loss. Over this period, the upper third of the watershed shifted from predominantly native grassland to an agricultural landscape, followed by widespread urbanization. The lower two-thirds of the watershed changed from a forested landscape to a mosaic of agriculture, urban, forest, and open water. Most channel length lost in the watershed over time was replaced by agriculture. Urban development gradually increased channel loss and disconnection from 1942 to 2011, particularly in the headwaters. Intensities of channel loss for both agriculture and urban increased over time. The two longest connected segments of channel loss came from the creation of two large impoundments, resulting in 46 km and 25 km of lost stream channel, respectively. Overall, the results from this study demonstrate that multiple and various land-use changes over long time periods can lead to rapid losses of large channel lengths as well as gradual (but increasing) losses of small channel lengths across all stream sizes. When these stream channel losses are taken into account, the environmental impacts of anthropogenic land-use change are compounded.