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.

Rising minimum temperatures contribute to 50 years of occupancy decline among cold-adapted Arctic and boreal butterflies in North America.

Global climate change has been identified as a potential driver of observed insect declines, yet in many regions, there are critical data gaps that make it difficult to assess how communities are responding to climate change. Poleward regions are of particular interest because warming is most rapid while biodiversity data are most sparse. Building on recent advances in occupancy modeling of presence-only data, we reconstructed 50 years (1970-2019) of butterfly occupancy trends in response to rising minimum temperatures in one of the most under-sampled regions of North America. Among 90 modeled species, we found that cold-adapted species are far more often in decline compared with their warm-adapted, more southernly distributed counterparts. Furthermore, in a post hoc analysis using species' traits, we find that species' range-wide average annual temperature is the only consistent predictor of occupancy changes. Species with warmer ranges were most likely to be increasing in occupancy. This trend results in the majority of butterflies increasing in occupancy probability over the last 50 years. Our results provide the first look at macroscale butterfly biodiversity shifts in high-latitude North America. These results highlight the potential of leveraging the wealth of presence-only data, the most abundant source of biodiversity data, for inferring changes in species distributions.

Ecological forecasts of insect range dynamics: a broad range of taxa includes winners and losers under future climate.

Species distribution models are the primary tools to project future species' distributions, but this complex task is influenced by data limitations and evolving best practices. The majority of the 53 studies we examined utilized correlative models and did not follow current best practices for validating retrospective or future environmental data layers. Despite this, a summary of results is largely unsurprising: shifts toward cooler regions, but otherwise mixed dynamics emphasizing winners and losers. Harmful insects were more likely to show positive outcomes compared with beneficial species. Our restricted ability to consider mechanisms complicates interpretation of any single study. To improve this area of modeling, more classic field and lab studies to uncover basic ecology and physiology are crucial.

How development and survival combine to determine the thermal sensitivity of insects.

Thermal performance curves (TPCs) depict variation in vital rates in response to temperature and have been an important tool to understand ecological and evolutionary constraints on the thermal sensitivity of ectotherms. TPCs allow for the calculation of indicators of thermal tolerance, such as minimum, optimum, and maximum temperatures that allow for a given metabolic function. However, these indicators are computed using only responses from surviving individuals, which can lead to underestimation of deleterious effects of thermal stress, particularly at high temperatures. Here, we advocate for an integrative framework for assessing thermal sensitivity, which combines both vital rates and survival probabilities, and focuses on the temperature interval that allows for population persistence. Using a collated data set of Lepidopteran development rate and survival measured on the same individuals, we show that development rate is generally limiting at low temperatures, while survival is limiting at high temperatures. We also uncover differences between life stages and across latitudes, with extended survival at lower temperatures in temperate regions. Our combined performance metric demonstrates similar thermal breadth in temperate and tropical individuals, an effect that only emerges from integration of both development and survival trends. We discuss the benefits of using this framework in future predictive and management contexts.

Population dynamics and drivers of the eastern monarch (Danaus plexippus) across its full annual cycle: a cross-scale synthesis of a model migratory species.

The monarch butterfly is arguably the best-known butterfly species throughout its global range. Declines in the size of the overwintering colonies in Mexico have sparked controversy regarding the conservation of the species and this controversy has been heightened since the United States Fish and Wildlife Service and International Union for the Conservation of Nature concluded that the eastern monarch populations were threatened (or in the case of United States Fish and Wildlife Service, warranted listing). Drivers of decline vary through space and time. Here, we present a synthesis of longitudinal monarch abundance studies that aim to disentangle the putative drivers of decline from one another. We find remarkable consistency that suggests monarch populations are indeed declining and that potential drivers of such decline shift over time. We strongly encourage future work on the species paired with mechanistic, experimental designs to address some long-standing knowledge gaps.

Consistent trait-temperature interactions drive butterfly phenology in both incidental and survey data.

Data availability limits phenological research at broad temporal and spatial extents. Butterflies are among the few taxa with broad-scale occurrence data, from both incidental reports and formal surveys. Incidental reports have biases that are challenging to address, but structured surveys are often limited seasonally and may not span full flight phenologies. Thus, how these data source compare in phenological analyses is unclear. We modeled butterfly phenology in relation to traits and climate using parallel analyses of incidental and survey data, to explore their shared utility and potential for analytical integration. One workflow aggregated "Pollard" surveys, where sites are visited multiple times per year; the other aggregated incidental data from online portals: iNaturalist and eButterfly. For 40 species, we estimated early (10%) and mid (50%) flight period metrics, and compared the spatiotemporal patterns and drivers of phenology across species and between datasets. For both datasets, inter-annual variability was best explained by temperature, and seasonal emergence was earlier for resident species overwintering at more advanced stages. Other traits related to habitat, feeding, dispersal, and voltinism had mixed or no impacts. Our results suggest that data integration can improve phenological research, and leveraging traits may predict phenology in poorly studied species.

LepTraits 1.0 A globally comprehensive dataset of butterfly traits.

Here, we present the largest, global dataset of Lepidopteran traits, focusing initially on butterflies (ca. 12,500 species records). These traits are derived from field guides, taxonomic treatments, and other literature resources. We present traits on wing size, phenology,voltinism, diapause/overwintering stage, hostplant associations, and habitat affinities (canopy, edge, moisture, and disturbance). This dataset will facilitate comparative research on butterfly ecology and evolution and our goal is to inspire future research collaboration and the continued development of this dataset.

Changes in climate drive recent monarch butterfly dynamics.

Declines in the abundance and diversity of insects pose a substantial threat to terrestrial ecosystems worldwide. Yet, identifying the causes of these declines has proved difficult, even for well-studied species like monarch butterflies, whose eastern North American population has decreased markedly over the last three decades. Three hypotheses have been proposed to explain the changes observed in the eastern monarch population: loss of milkweed host plants from increased herbicide use, mortality during autumn migration and/or early-winter resettlement and changes in breeding-season climate. Here, we use a hierarchical modelling approach, combining data from >18,000 systematic surveys to evaluate support for each of these hypotheses over a 25-yr period. Between 2004 and 2018, breeding-season weather was nearly seven times more important than other factors in explaining variation in summer population size, which was positively associated with the size of the subsequent overwintering population. Although data limitations prevent definitive evaluation of the factors governing population size between 1994 and 2003 (the period of the steepest monarch decline coinciding with a widespread increase in herbicide use), breeding-season weather was similarly identified as an important driver of monarch population size. If observed changes in spring and summer climate continue, portions of the current breeding range may become inhospitable for monarchs. Our results highlight the increasingly important contribution of a changing climate to insect declines.

Ecological mechanism of climate-mediated selection in a rapidly evolving invasive species.

Recurring seasonal changes can lead to the evolution of phenological cues. For example, many arthropods undergo photoperiodic diapause, a programmed developmental arrest induced by short autumnal day length. The selective mechanisms that determine the timing of autumnal diapause initiation have not been empirically identified. We quantified latitudinal clines in genetically determined diapause timing of an invasive mosquito, Aedes albopictus, on two continents. We show that variation in diapause timing within and between continents is explained by a novel application of a growing degree day (GDD) model that delineates a location-specific deadline after which it is not possible to complete an additional full life cycle. GDD models are widely used to predict spring phenology by modelling growth and development as physiological responses to ambient temperatures. Our results show that the energy accumulation dynamics represented by GDD models have also led to the evolution of an anticipatory life-history cue in autumn.

A novel curation system to facilitate data integration across regional citizen science survey programs.

Integrative modeling methods can now enable macrosystem-level understandings of biodiversity patterns, such as range changes resulting from shifts in climate or land use, by aggregating species-level data across multiple monitoring sources. This requires ensuring that taxon interpretations match up across different sources. While encouraging checklist standardization is certainly an option, coercing programs to change species lists they have used consistently for decades is rarely successful. Here we demonstrate a novel approach for tracking equivalent names and concepts, applied to a network of 10 regional programs that use the same protocols (so-called "Pollard walks") to monitor butterflies across America north of Mexico. Our system involves, for each monitoring program, associating the taxonomic authority (in this case one of three North American butterfly fauna treatments: Pelham, 2014; North American Butterfly Association, Inc., 2016; Opler & Warren, 2003) that shares the most similar overall taxonomic interpretation to the program's working species list. This allows us to define each term on each program's list in the context of the appropriate authority's species concept and curate the term alongside its authoritative concept. We then aligned the names representing equivalent taxonomic concepts among the three authorities. These stepping stones allow us to bridge a species concept from one program's species list to the name of the equivalent in any other program, through the intermediary scaffolding of aligned authoritative taxon concepts. Using a software tool we developed to access our curation system, a user can link equivalent species concepts between data collecting agencies with no specialized knowledge of taxonomic complexities.

Matching expert range maps with species distribution model predictions.

Species' range maps based on expert opinion are a critical resource for conservation planning. Expert maps are usually accompanied by species descriptions that specify sources of internal range heterogeneity, such as habitat associations, but these are rarely considered when using expert maps for analyses. We developed a quantitative metric (expert score) to evaluate the agreement between an expert map and a habitat probability surface obtained from a species distribution model. This method rewards both the avoidance of unsuitable sites and the inclusion of suitable sites in the expert map. We obtained expert maps of 330 butterfly species from each of 2 widely used North American sources (Glassberg [1999, 2001] and Scott [1986]) and computed species-wise expert scores for each. Overall, the Glassberg maps secured higher expert scores than Scott (0.61 and 0.41, respectively) due to the specific rules (e.g., Glassberg only included regions where the species was known to reproduce whereas Scott included all areas a species expanded to each year) they used to include or exclude areas from ranges. The predictive performance of expert maps was almost always hampered by the inclusion of unsuitable sites, rather than by exclusion of suitable sites (deviance outside of expert maps was extremely low). Map topology was the primary predictor of expert performance rather than any factor related to species characteristics such as mobility. Given the heterogeneity and discontinuity of suitable landscapes, expert maps drawn with more detail are more likely to agree with species distribution models and thus minimize both commission and omission errors.

Concordancia entre los Mapas de Extensión Realizados por Expertos y las Predicciones de los Modelos de Distribución de Especies Resumen Los mapas de extensión de especies basados en la opinión de expertos son un recurso de suma importancia para la planeación de la conservación. Los mapas realizados por expertos generalmente van acompañados de las descripciones de las especies que detallan el origen de la heterogeneidad interna de la distribución, como las asociaciones entre hábitats, pero rara vez se consideran estas descripciones cuando se usan los mapas de expertos para un análisis. Desarrollamos una medida cuantitativa (puntaje de expertos) para evaluar la concordancia entre un mapa realizado por expertos y una superficie probable de hábitat obtenida a partir del modelo de distribución de especies (SDM). Este método recompensa tanto a la evasión de sitios inadecuados como a la inclusión de sitios adecuados en el mapa realizado por expertos. Obtuvimos los mapas realizados por expertos para 330 especies de mariposas a partir de dos fuentes norteamericanas usadas ampliamente (Glassberg [1999, 2001] y Scott [1986]) y calculamos los puntajes de expertos, hablando de cada especie, para cada mapa. En general, los mapas de Glassberg aseguraron puntajes de expertos más altos que los de Scott (0.61 y 0.41 respectivamente) debido a las reglas específicas (p. ej., Glassberg sólo incluyó las regiones en donde es sabido que la especie se reproduce, mientras que Scott incluyó todas las áreas a las que la especie se expandió cada año) que cada una usa para incluir o excluir áreas de las distribuciones. El desempeño pronosticado de los mapas realizados por expertos casi siempre se vio afectado por la inclusión de los sitios inadecuados, en lugar de estar afectado por la exclusión de sitios adecuados (la desviación fuera de los mapas realizados por expertos fue extremadamente baja). La topología del mapa fue el indicador primario del desempeño de los expertos en lugar de cualquier factor relacionado con las características de la especie, como la movilidad. Dada la heterogeneidad y la discontinuidad de los paisajes adecuados, los mapas realizados por expertos dibujados con mayor detalle tienen una mayor probabilidad de concordar con los SMD y por lo tanto minimizar los errores de comisión y de omisión.

Butterfly abundance declines over 20 years of systematic monitoring in Ohio, USA.

Severe insect declines make headlines, but they are rarely based on systematic monitoring outside of Europe. We estimate the rate of change in total butterfly abundance and the population trends for 81 species using 21 years of systematic monitoring in Ohio, USA. Total abundance is declining at 2% per year, resulting in a cumulative 33% reduction in butterfly abundance. Three times as many species have negative population trends compared to positive trends. The rate of total decline and the proportion of species in decline mirror those documented in three comparable long-term European monitoring programs. Multiple environmental changes such as climate change, habitat degradation, and agricultural practices may contribute to these declines in Ohio and shift the makeup of the butterfly community by benefiting some species over others. Our analysis of life-history traits associated with population trends shows an impact of climate change, as species with northern distributions and fewer annual generations declined more rapidly. However, even common and invasive species associated with human-dominated landscapes are declining, suggesting widespread environmental causes for these trends. Declines in common species, although they may not be close to extinction, will have an outsized impact on the ecosystem services provided by insects. These results from the most extensive, systematic insect monitoring program in North America demonstrate an ongoing defaunation in butterflies that on an annual scale might be imperceptible, but cumulatively has reduced butterfly numbers by a third over 20 years.

Habitat edge responses of generalist predators are predicted by prey and structural resources.

Generalist predators are thought to be less vulnerable to habitat fragmentation because they use diverse resources across larger spatial scales than specialist predators. Thus, it has been suggested that generalist predators may respond positively to habitat edges or demonstrate no edge response, because they can potentially use prey resources equally well on both sides of the habitat edge. However, most predictions about generalist predator responses to the habitat edge are based solely on prey resources, without consideration of other potential drivers. For instance, structural resources are essential for some species to build webs to capture prey or to avoid intraguild predation and cannibalism. In this study, we used both prey and structural resources to predict the response of four predator functional groups (hunting spiders, web-building spiders, aerial predators, and epigeic predators that feed on the detrital/algal food web) to a habitat edge between two salt-marsh grasses (Spartina alterniflora and Spartina patens). We found that generalist predators largely demonstrated negative responses to the habitat edge and had distinct habitat associations. Positive edge responses were only observed in one functional group (hunting spiders), and this pattern was driven by the two most abundant species. Negative responses to the habitat edge were more common among taxa and were better explained by structural resources rather than prey resources in the two habitats. Although it is generally acknowledged that specialists decline in fragmented habitats, generalists are thought to be more resilient. However, our research demonstrates that even generalists have habitat structural or food resource requirements that may limit their resilience to habitat loss and fragmentation.

Multiscale seasonal factors drive the size of winter monarch colonies.

Monarch butterflies in eastern North America have declined by 84% on Mexican wintering grounds since the observed peak in 1996. However, coarse-scale population indices from northern US breeding grounds do not show a consistent downward trend. This discrepancy has led to speculation that autumn migration may be a critical limiting period. We address this hypothesis by examining the role of multiscale processes impacting monarchs during autumn, assessed using arrival abundances at all known winter colony sites over a 12-y period (2004-2015). We quantified effects of continental-scale (climate, landscape greenness, and disease) and local-scale (colony habitat quality) drivers of spatiotemporal trends in winter colony sizes. We also included effects of peak summer and migratory population indices. Our results demonstrate that higher summer abundance on northern breeding grounds led to larger winter colonies as did greener autumns, a proxy for increased nectar availability in southern US floral corridors. Colony sizes were also positively correlated with the amount of local dense forest cover and whether they were located within the Monarch Butterfly Biosphere Reserve, but were not influenced by disease rates. Although we demonstrate a demographic link between summer and fine-scale winter population sizes, we also reveal that conditions experienced during, and at the culmination of, autumn migration impact annual dynamics. Monarchs face a growing threat if floral resources and winter habitat availability diminish under climate change. Our study tackles a long-standing gap in the monarch's annual cycle and highlights the importance of evaluating migratory conditions to understand mechanisms governing long-term population trends.

Complex community and evolutionary responses to habitat fragmentation and habitat edges: what can we learn from insect science?

Habitat fragmentation is the primary factor leading to species extinction worldwide and understanding how species respond to habitat edges is critical for understanding the effects of fragmentation on insect diversity in both natural and managed landscapes. Most studies on insect responses to the habitat edge focus on bottom-up changes in resources. Only a few recent studies have examined multi-trophic responses to habitat edges; the results of these studies highlight the problem that we lack a conceptual framework to understand the complex results observed when a single species' response to an edge 'cascades' throughout the food web in ways that are currently not predictable. Recent research from insect systems suggests that habitat edge responses cascade both up and down multi-trophic foodwebs and these altered species interactions may affect evolutionary processes. Future studies that investigate the effects of habitat edges on both ecological and evolutionary dynamics can help to fill these knowledge gaps and we suggest that insects, with short generation times, present an ideal opportunity to do so.

Multitaxonomic diversity patterns along a desert riparian-upland gradient.

Riparian areas are noted for their high biodiversity, but this has rarely been tested across a wide range of taxonomic groups. We set out to describe species richness, species abundance, and community similarity patterns for 11 taxonomic groups (forbs & grasses, shrubs, trees, solpugids, spiders, scarab beetles, butterflies, lizards, birds, rodents, and mammalian carnivores) individually and for all groups combined along a riparian-upland gradient in semiarid southeastern Arizona, USA. Additionally, we assessed whether biological characteristics could explain variation in diversity along the gradient using five traits (trophic level, body size, life span, thermoregulatory mechanism, and taxonomic affiliation). At the level of individual groups diversity patterns varied along the gradient, with some having greater richness and/or abundance in riparian zones whereas others were more diverse and/or abundant in upland zones. Across all taxa combined, riparian zones contained significantly more species than the uplands. Community similarity between riparian and upland zones was low, and beta diversity was significantly greater than expected for most taxonomic groups, though biological traits explained little variance in diversity along the gradient. These results indicate heterogeneity amongst taxa in how they respond to the factors that structure ecological communities in riparian landscapes. Nevertheless, across taxonomic groups the overall pattern is one of greater species richness and abundance in riparian zones, coupled with a distinct suite of species.

Tracking climate impacts on the migratory monarch butterfly.

Understanding the impacts of climate on migratory species is complicated by the fact that these species travel through several climates that may be changing in diverse ways throughout their complete migratory cycle. Most studies are not designed to tease out the direct and indirect effects of climate at various stages along the migration route. We assess the impacts of spring and summer climate conditions on breeding monarch butterflies, a species that completes its annual migration cycle over several generations. No single, broad-scale climate metric can explain summer breeding phenology or the substantial year-to-year fluctuations observed in population abundances. As such, we built a Poisson regression model to help explain annual arrival times and abundances in the Midwestern United States. We incorporated the climate conditions experienced both during a spring migration/breeding phase in Texas as well as during subsequent arrival and breeding during the main recruitment period in Ohio. Using data from a state-wide butterfly monitoring network in Ohio, our results suggest that climate acts in conflicting ways during the spring and summer seasons. High spring precipitation in Texas is associated with the largest annual population growth in Ohio and the earliest arrival to the summer breeding ground, as are intermediate spring temperatures in Texas. On the other hand, the timing of monarch arrivals to the summer breeding grounds is not affected by climate conditions within Ohio. Once in Ohio for summer breeding, precipitation has minimal impacts on overall abundances, whereas warmer summer temperatures are generally associated with the highest expected abundances, yet this effect is mitigated by the average seasonal temperature of each location in that the warmest sites receive no benefit of above average summer temperatures. Our results highlight the complex relationship between climate and performance for a migrating species and suggest that attempts to understand how monarchs will be affected by future climate conditions will be challenging.

Do edge responses cascade up or down a multi-trophic food web?

Despite nearly 100 years of edge studies, there has been little effort to document how edge responses 'cascade' to impact multi-trophic food webs. We examined changes within two, four-tiered food webs located on opposite sides of a habitat edge. Based on a 'bottom-up' resource-based model, we predicted plant resources would decline near edges, causing similar declines in specialist herbivores and their associated predators, while a generalist predator was predicted to increase due to complementary resource use. As predicted, we found declines in both specialist herbivores and predators near edges, but, contrary to expectations, this was not driven by gradients in plant resources. Instead, the increase in generalist predators near edges offers one alternative explanation for the observed declines. Furthermore, our results suggest how recent advances in food web theory could improve resource-based edge models, and vice versa.

Avian predators are less abundant during periodical cicada emergences, but why?

Despite a substantial resource pulse, numerous avian insectivores known to depredate periodical cicadas (Magicicada spp.) are detected less commonly during emergence years than in either the previous or following years. We used data on periodical cicada calls collected by volunteers conducting North American Breeding Bird Surveys within the range of cicada Brood X to test three hypotheses for this observation: lower detection rates could be caused by bird calls being obscured by cicada calls ("detectability" hypothesis), by birds avoiding areas with cicadas ("repel" hypothesis), or because bird abundances are generally lower during emergence years for some reason unrelated to the current emergence event ("true decline" hypothesis). We tested these hypotheses by comparing bird detections at stations coincident with calling cicadas vs. those without calling cicadas in the year prior to and during cicada emergences. At four distinct levels (stop, route, range, and season), parallel declines of birds in groups exposed and not exposed to cicada calls supported the true decline hypothesis. We discuss several potential mechanisms for this pattern, including the possibility that it is a consequence of the ecological and evolutionary interactions between predators of this extraordinary group of insects.