Cross-Site Comparisons of Dryland Ecosystem Response to Climate Change in the US Long-Term Ecological Research Network.

Long-term observations and experiments in diverse drylands reveal how ecosystems and services are responding to climate change. To develop generalities about climate change impacts at dryland sites, we compared broadscale patterns in climate and synthesized primary production responses among the eight terrestrial, nonforested sites of the United States Long-Term Ecological Research (US LTER) Network located in temperate (Southwest and Midwest) and polar (Arctic and Antarctic) regions. All sites experienced warming in recent decades, whereas drought varied regionally with multidecadal phases. Multiple years of wet or dry conditions had larger effects than single years on primary production. Droughts, floods, and wildfires altered resource availability and restructured plant communities, with greater impacts on primary production than warming alone. During severe regional droughts, air pollution from wildfire and dust events peaked. Studies at US LTER drylands over more than 40 years demonstrate reciprocal links and feedbacks among dryland ecosystems, climate-driven disturbance events, and climate change.

How long do population level field experiments need to be? Utilising data from the 40-year-old LTER network.

We utilise the wealth of data accessible through the 40-year-old Long-Term Ecological Research (LTER) network to ask if aspects of the study environment or taxa alter the duration of research necessary to detect consistent results. To do this, we use a moving-window algorithm. We limit our analysis to long-term (> 10 year) press experiments recording organismal abundance. We find that studies conducted in dynamic abiotic environments need longer periods of study to reach consistent results, as compared to those conducted in more moderated environments. Studies of plants were more often characterised by spurious results than those on animals. Nearly half of the studies we investigated required 10 years or longer to become consistent, where all significant trends agreed in direction, and four studies (of 100) required longer than 20 years. Here, we champion the importance of long-term data and bolster the value of multi-decadal experiments in understanding, explaining and predicting long-term trends.

Bioenergy landscapes drive trophic shifts in generalist ants.

Changes in trophic niche-the pathways through which an organism obtains energy and nutrients-are a fundamental way in which organisms respond to environmental conditions. But the capacity for species to alter their trophic niches in response to global change, and the ways they do so when able, remain largely unknown. Here we examine food webs in three long-term and large-scale experiments to test how resource availability and nutritional requirements interact to determine an organism's trophic niche in the context of one of the largest global trends in land use-the rise in bioenergy production. We use carbon and nitrogen stable isotope analyses to characterize arthropod food webs across three biofuel crops representing a gradient in plant resource richness (corn monocultures, fields dominated by native switchgrass and restored prairie), and to quantify changes in the trophic niche of a widespread generalist ant species across habitats. In doing so, we measure the effects of basal resource richness on food chain length, niche breadth and trophic position. We frame our results in the context of two hypotheses that explain variation in trophic niche-the niche variation hypothesis which emphasizes the importance of resource diversity and ecological opportunity, and the optimal diet hypothesis which emphasizes dietary constraints and the availability of optimal resources. Increasing plant richness lengthened food chains by 10%-20% compared to monocultures. Niche breadths of generalist ants did not vary with resource richness, suggesting they were limited by optimal diet requirements and constraints rather than by ecological opportunity. The ants instead responded to changes in plant richness by shifting their estimated trophic position. In resource-poor monocultures, the ants were top predators, sharing a trophic position with predatory spiders. In resource-rich environments, in contrast, the ants were omnivores, relying on a mix of animal prey and plant-based resources. In addition to highlighting novel ecosystem impacts of alternate bioenergy landscapes, our results suggest that niche breadth and trophic diversification depend more on the presence of optimal resources than on ecological opportunity alone.

Maintaining historic disturbance regimes increases species' resilience to catastrophic hurricanes.

As habitat loss and fragmentation, urbanization, and global climate change accelerate, conservation of rare ecosystems increasingly relies on human intervention. However, any conservation strategy is vulnerable to unpredictable, catastrophic events. Whether active management increases or decreases a system's resilience to these events remains unknown. Following Hurricane Irma's landfall in our habitat restoration study sites, we found that rare ecosystems with active, human-imposed management suffered less damage in a hurricane's path than unmanaged systems. At the center of Irma's landfall, we found Croton linearis' (a locally rare plant that is the sole host for two endangered butterfly species) survival and population growth rates in the year of the hurricane were higher in previously managed plots than in un-managed controls. In the periphery of Irma's circulation, the effect of prior management was stronger than that of the hurricane. Maintaining the historical disturbance regime thus increased the resilience of the population to major hurricane disturbance. As climate change increases the probability and intensity of severe hurricanes, human management of disturbance-adapted landscapes will become increasingly important for maintaining populations of threatened species in a storm's path. Doing nothing will accelerate extinction.

Long-term research avoids spurious and misleading trends in sustainability attributes of no-till.

Agricultural management recommendations based on short-term studies can produce findings inconsistent with long-term reality. Here, we test the long-term environmental sustainability and profitability of continuous no-till agriculture on yield, soil water availability, and N2 O fluxes. Using a moving window approach, we investigate the development and stability of several attributes of continuous no-till as compared to conventional till agriculture over a 29-year period at a site in the upper Midwest, US. Over a decade is needed to detect the consistent effects of no-till. Both crop yield and soil water availability required 15 years or longer to generate patterns consistent with 29-year trends. Only marginal trends for N2 O fluxes appeared in this period. Relative profitability analysis suggests that after initial implementation, 86% of periods between 10 and 29 years recuperated the initial expense of no-till implementation, with the probability of higher relative profit increasing with longevity. Importantly, statistically significant but misleading short-term trends appeared in more than 20% of the periods examined. Results underscore the importance of decadal and longer studies for revealing consistent dynamics and emergent outcomes of no-till agriculture, shown to be beneficial in the long term.

Movement and Demography of At-Risk Butterflies: Building Blocks for Conservation.

The number of insect species at risk of population decline and extinction is increasing rapidly. Yet we know almost nothing about the ecology of these species, except for at-risk butterflies. A growing body of literature shows how butterfly vital rates, including demography and movement, are essential for guiding conservation and recovery. History has shown us that without these data, conservation decisions often weaken, rather than enhance, population viability. This is especially true in changing landscapes. We review knowledge of vital rates across all at-risk butterflies. We have information on movement for 17 of 283 butterfly species and information on demography for 19 species. We find that habitat-specific movement behavior is key to understanding how to connect populations, and habitat-specific demography is central to managing habitats. Methods and analyses worked out for butterflies can provide a scaffold around which to build studies for the conservation of other at-risk insects.

Landscape heterogeneity is key to forecasting outcomes of plant reintroduction.

Conservation and restoration projects often involve starting new populations by introducing individuals into portions of their native or projected range. Such efforts can help meet many related goals, including habitat creation, ecosystem service provisioning, assisted migration, and the reintroduction of imperiled species following local extirpation. The outcomes of reintroduction efforts, however, are highly variable, with results ranging from local extinction to dramatic population growth; reasons for this variation remain unclear. Here, we ask whether population growth following plant reintroductions is governed by variation at two scales: the scale of individual habitat patches to which individuals are reintroduced, and larger among-landscape scales in which similar patches may be situated in landscapes that differ in matrix type, soil conditions, and other factors. Quantifying demographic variation at these two scales will help prioritize locations for introduction and, once introductions take place, forecast population growth. This work took place within a large-scale habitat fragmentation experiment, where individuals of two perennial forb species were reintroduced into eight replicate ~50-ha landscapes, each containing a set of five ~1-ha patches that varied in their degree of isolation (connected by habitat corridors or unconnected) and edge-to-area ratio. Using data on individual growth, survival, reproductive output, and recruitment collected one to two years after reintroduction, we developed models to forecast population growth, then compared forecasts to observed population sizes, three and six years later. Both the type of patch (connected and unconnected) and identity of the landscape to which individuals were reintroduced had effects on forecasted population growth rates, but only variation associated with landscape identity was an accurate predictor of subsequently observed population growth rates. Models that did not include landscape identity had minimal forecasting ability, revealing the key importance of variation at this scale for accurate prediction. Of the five demographic rates used to model population dynamics, seed production was the most important source of forecast error in population growth rates. Our results point to the importance of accounting for landscape-scale variation in demographic models and demonstrate how such models might assist with prioritizing particular landscapes for species reintroduction projects.

Connectivity increases trophic subsidies in fragmented landscapes.

Landscape corridors mitigate the negative effects of habitat fragmentation by increasing dispersal. Corridors also increase biodiversity in connected habitat fragments, suggestive of metacommunity dynamics. What is unknown in this case is the mechanisms through which metacommunity dynamics act. Working in a large-scale fragmentation experiment, we tested the effect of corridors on the movement of prey species and subsequent effects on predator nutrition (which we call trophic subsidies). We enriched plants of central patches with 15 N, then measured δ15 N in green lynx spiders, the most abundant insect predator, in patches that were either connected to or isolated from the enriched patch. We found that corridors increased prey movement, as they increased spider δ15 N by 40% in connected patches. Corridors also improved spider body condition, increasing nitrogen relative to carbon. We suggest a novel mechanism, trophic subsidies, through which corridors may increase the stability or size of populations in connected landscapes.

How fragmentation and corridors affect wind dynamics and seed dispersal in open habitats.

Determining how widespread human-induced changes such as habitat loss, landscape fragmentation, and climate instability affect populations, communities, and ecosystems is one of the most pressing environmental challenges. Critical to this challenge is understanding how these changes are affecting the movement abilities and dispersal trajectories of organisms and what role conservation planning can play in promoting movement among remaining fragments of suitable habitat. Whereas evidence is mounting for how conservation strategies such as corridors impact animal movement, virtually nothing is known for species dispersed by wind, which are often mistakenly assumed to not be limited by dispersal. Here, we combine mechanistic dispersal models, wind measurements, and seed releases in a large-scale experimental landscape to show that habitat corridors affect wind dynamics and seed dispersal by redirecting and bellowing airflow and by increasing the likelihood of seed uplift. Wind direction interacts with landscape orientation to determine when corridors provide connectivity. Our results predict positive impacts of connectivity and patch shape on species richness of wind-dispersed plants, which we empirically illustrate using 12 y of data from our experimental landscapes. We conclude that habitat fragmentation and corridors strongly impact the movement of wind-dispersed species, which has community-level consequences.

Disentangling fragmentation effects on herbivory in understory plants of longleaf pine savanna.

Habitat fragmentation affects species and their interactions through intertwined mechanisms that include changes to fragment area, shape, connectivity and distance to edge. Disentangling these pathways is a fundamental challenge of landscape ecology and will help identify ecological processes important for management of rare species or restoration of fragmented habitats. In a landscape experiment that manipulated connectivity, fragment shape, and distance to edge while holding fragment area constant, we examined how fragmentation impacts herbivory and growth of nine plant species in longleaf pine savanna. Probability of herbivory in open habitat was strongly dependent on proximity to forest edge for every species, increasing with distance to edge in six species (primarily grasses and annual forbs) and decreasing in three species (perennial forbs and a shrub). In the two species of perennial forbs, these edge effects were dependent on fragment shape; herbivory strongly decreased with distance to edge in fragments of two shapes, but not in a third shape. For most species, however, probability of herbivory was unrelated to connectivity or fragment shape. Growth was generally determined more strongly by leaf herbivory than by distance to edge, fragment shape, or connectivity. Taken together, these results demonstrate consistently strong edge effects on herbivory, one of the most important biotic factors determining plant growth and demography. Our results contrast with the generally inconsistent results of observational studies, likely because our experimental approach enabled us to tease apart landscape processes that are typically confounded.

Connectivity from a different perspective: comparing seed dispersal kernels in connected vs. unfragmented landscapes.

Habitat fragmentation can create significant impediments to dispersal. A technique to increase dispersal between otherwise isolated fragments is the use of corridors. Although previous studies have compared dispersal between connected fragments to dispersal between unconnected fragments, it remains unknown how dispersal between fragments connected by a corridor compares to dispersal in unfragmented landscapes. To assess the extent to which corridors can restore dispersal in fragmented landscapes to levels observed in unfragmented landscapes, we employed a stable-isotope marking technique to track seeds within four unfragmented landscapes and eight experimental landscapes with fragments connected by corridors. We studied two wind- and two bird-dispersed plant species, because previous community-based research showed that dispersal mode explains how connectivity effects vary among species. We constructed dispersal kernels for these species in unfragmented landscapes and connected fragments by marking seeds in the center of each landscape with 'IN and then recovering marked seeds in seed traps at distances up to 200 m. For the two wind-dispersed plants, seed dispersal kernels were similar in unfragmented landscapes and connected fragments. In contrast, dispersal kernels of bird-dispersed seeds were both affected by fragmentation and differed in the direction of the impact: Morella cerifera experienced more and Rhus copallina experienced less long-distance dispersal in unfragmented than in connected landscapes. These results show that corridors can facilitate dispersal probabilities comparable to those observed in unfragmented landscapes. Although dispersal mode may provide useful broad predictions, we acknowledge that similar species may respond uniquely due to factors such as seasonality and disperser behavior. Our results further indicate that prior work has likely underestimated dispersal distances of wind-dispersed plants and that factors altering long-distance dispersal may have a greater impact on the spread of species than previously thought.

Drosophila suzukii: the genetic footprint of a recent, worldwide invasion.

Native to Asia, the soft-skinned fruit pest Drosophila suzukii has recently invaded the United States and Europe. The eastern United States represents the most recent expansion of their range, and presents an opportunity to test alternative models of colonization history. Here, we investigate the genetic population structure of this invasive fruit fly, with a focus on the eastern United States. We sequenced six X-linked gene fragments from 246 individuals collected from a total of 12 populations. We examine patterns of genetic diversity within and between populations and explore alternative colonization scenarios using approximate Bayesian computation. Our results indicate high levels of nucleotide diversity in this species and suggest that the recent invasions of Europe and the continental United States are independent demographic events. More broadly speaking, our results highlight the importance of integrating population structure into demographic models, particularly when attempting to reconstruct invasion histories. Finally, our simulation results illustrate the general challenge in reconstructing invasion histories using genetic data and suggest that genome-level data are often required to distinguish among alternative demographic scenarios.

The influence of habitat fragmentation on multiple plant-animal interactions and plant reproduction.

Despite broad recognition that habitat loss represents the greatest threat to the world's biodiyersity, a mechanistic understanding of how habitat loss and associated fragmentation affect ecological systems has proven remarkably challenging. The challenge stems from the multiple interdependent ways that landscapes change following fragmentation and the ensuing complex impacts on populations and communities of interacting species. We confronted these challenges by evaluating how fragmentation affects individual plants through interactions with animals, across five herbaceous species native to longleaf pine savannas. We created a replicated landscape experiment that provides controlled tests of three major fragmentation effects (patch isolation, patch shape [i.e., edge-to-area ratio], and distance to edge), established experimental founder populations of the five species to control for spatial distributions and densities of individual plants, and employed structural equation modeling to evaluate the effects of fragmentation on plant reproductive output and the degree to which these impacts are mediated through altered herbivory, pollination, or pre-dispersal seed predation. Across species, the most consistent response to fragmentation was a reduction in herbivory. Herbivory, however, had little impact.on plant reproductive output, and thus we found little evidence for any resulting benefit to plants in fragments. In contrast, fragmentation rarely impacted pollination or pre-dispersal seed predation, but both of these interactions had strong and consistent impacts on plant reproductive output. As a result, our models robustly predicted plant reproductive output (r2 = 0.52-0.70), yet due to the weak effects of fragmentation on pollination and pre-dispersal seed predation, coupled with the weak effect of herbivory on plant reproduction, the effects of fragmentation on reproductive output were generally small in magnitude and inconsistent. This work provides mechanistic insight into landscape-scale variation in plant reproductive success, the relative importance of plant-animal interactions for structuring these dynamics, and the nuanced nature of how habitat fragmentation can affect populations and communities of interacting species.

Landscape corridors can increase invasion by an exotic species and reduce diversity of native species.

Landscape corridors are commonly used to mitigate negative effects of habitat fragmentation, but concerns persist that they may facilitate the spread of invasive species. In a replicated landscape experiment of open habitat, we measured effects of corridors on the invasive fire ant, Solenopsis invicta, and native ants. Fire ants have two social forms: polygyne, which tend to disperse poorly but establish at high densities, and monogyne, which disperse widely but establish at lower densities. In landscapes dominated by polygyne fire ants, fire ant abundance was higher and native ant diversity was lower in habitat patches connected by corridors than in unconnected patches. Conversely, in landscapes dominated by monogyne fire ants, connectivity had no influence on fire ant abundance and native ant diversity. Polygyne fire ants dominated recently created landscapes, suggesting that these corridor effects may be transient. Our results suggest that corridors can facilitate invasion and they highlight the importance of considering species' traits when assessing corridor utility.

Defining and evaluating the umbrella species concept for conserving and restoring landscape connectivity.

Conserving or restoring landscape connectivity between patches of breeding habitat is a common strategy to protect threatened species from habitat fragmentation. By managing connectivity for some species, usually charismatic vertebrates, it is often assumed that these species will serve as conservation umbrellas for other species. We tested this assumption by developing a quantitative method to measure overlap in dispersal habitat of 3 threatened species-a bird (the umbrella), a butterfly, and a frog-inhabiting the same fragmented landscape. Dispersal habitat was determined with Circuitscape, which was parameterized with movement data collected for each species. Despite differences in natural history and breeding habitat, we found substantial overlap in the spatial distributions of areas important for dispersal of this suite of taxa. However, the intuitive umbrella species (the bird) did not have the highest overlap with other species in terms of the areas that supported connectivity. Nevertheless, we contend that when there are no irreconcilable differences between the dispersal habitats of species that cohabitate on the landscape, managing for umbrella species can help conserve or restore connectivity simultaneously for multiple threatened species with different habitat requirements.

Potential negative ecological effects of corridors.

Despite many studies showing that landscape corridors increase dispersal and species richness for disparate taxa, concerns persist that corridors can have unintended negative effects. In particular, some of the same mechanisms that underlie positive effects of corridors on species of conservation interest may also increase the spread and impact of antagonistic species (e.g., predators and pathogens), foster negative effects of edges, increase invasion by exotic species, increase the spread of unwanted disturbances such as fire, or increase population synchrony and thus reduce persistence. We conducted a literature review and meta-analysis to evaluate the prevalence of each of these negative effects. We found no evidence that corridors increase unwanted disturbance or non-native species invasion; however, these have not been well-studied concerns (1 and 6 studies, respectively). Other effects of corridors were more often studied and yielded inconsistent results; mean effect sizes were indistinguishable from zero. The effect of edges on abundances of target species was as likely to be positive as negative. Corridors were as likely to have no effect on antagonists or population synchrony as they were to increase those negative effects. We found 3 deficiencies in the literature. First, despite studies on how corridors affect predators, there are few studies of related consequences for prey population size and persistence. Second, properly designed studies of negative corridor effects are needed in natural corridors at scales larger than those achievable in experimental systems. Third, studies are needed to test more targeted hypotheses about when corridor-mediated effects on invasive species or disturbance may be negative for species of management concern. Overall, we found no overarching support for concerns that construction and maintenance of habitat corridors may result in unintended negative consequences. Negative edge effects may be mitigated by widening corridors or softening edges between corridors and the matrix. Other negative effects are relatively small and manageable compared with the large positive effects of facilitating dispersal and increasing diversity of native species.

Insecticides, more than herbicides, land use, and climate, are associated with declines in butterfly species richness and abundance in the American Midwest.

Mounting evidence shows overall insect abundances are in decline globally. Habitat loss, climate change, and pesticides have all been implicated, but their relative effects have never been evaluated in a comprehensive large-scale study. We harmonized 17 years of land use, climate, multiple classes of pesticides, and butterfly survey data across 81 counties in five states in the US Midwest. We find community-wide declines in total butterfly abundance and species richness to be most strongly associated with insecticides in general, and for butterfly species richness the use of neonicotinoid-treated seeds in particular. This included the abundance of the migratory monarch (Danaus plexippus), whose decline is the focus of intensive debate and public concern. Insect declines cannot be understood without comprehensive data on all putative drivers, and the 2015 cessation of neonicotinoid data releases in the US will impede future research.

Diversity of plant evolutionary lineages promotes arthropod diversity.

Large-scale habitat destruction and climate change result in the non-random loss of evolutionary lineages, reducing the amount of evolutionary history represented in ecological communities. Yet, we have limited understanding of the consequences of evolutionary history on the structure of food webs and the services provided by biological communities. Drawing on 11 years of data from a long-term plant diversity experiment, we show that evolutionary history of plant communities - measured as phylogenetic diversity - strongly predicts diversity and abundance of herbivorous and predatory arthropods. Effects of plant species richness on arthropods become stronger when phylogenetic diversity is high. Plant phylogenetic diversity explains predator and parasitoid richness as strongly as it does herbivore richness. Our findings indicate that accounting for evolutionary relationships is critical to understanding the severity of species loss for food webs and ecosystems, and for developing conservation and restoration policies.

How complex do models need to be to predict dispersal of threatened species through matrix habitats?

Persistence of species in fragmented landscapes depends on dispersal among suitable breeding sites, and dispersal is often influenced by the "matrix" habitats that lie between breeding sites. However, measuring effects of different matrix habitats on movement and incorporating those differences into spatially explicit models to predict dispersal is costly in terms of time and financial resources. Hence a key question for conservation managers is: Do more costly, complex movement models yield more accurate dispersal predictions? We compared the abilities of a range of movement models, from simple to complex, to predict the dispersal of an endangered butterfly, the Saint Francis' satyr (Neonympha mitchellii francisci). The value of more complex models differed depending on how value was assessed. Although the most complex model, based on detailed movement behaviors, best predicted observed dispersal rates, it was only slightly better than the simplest model, which was based solely on distance between sites. Consequently, a parsimony approach using information criteria favors the simplest model we examined. However, when we applied the models to a larger landscape that included proposed habitat restoration sites, in which the composition of the matrix was different than the matrix surrounding extant breeding sites, the simplest model failed to identify a potentially important dispersal barrier, open habitat that butterflies rarely enter, which may completely isolate some of the proposed restoration sites from other breeding sites. Finally, we found that, although the gain in predicting dispersal with increasing model complexity was small, so was the increase in financial cost. Furthermore, a greater fit continued to accrue with greater financial cost, and more complex models made substantially different predictions than simple models when applied to a novel landscape in which butterflies are to be reintroduced to bolster their populations. This suggests that more complex models might be justifiable on financial grounds. Our results caution against a pure parsimony approach to deciding how complex movement models need to be to accurately predict dispersal through the matrix, especially if the models are to be applied to novel or modified landscapes.