Pollinator activity and flowering in agricultural weeds in Sweden.

The extent to which weeds in arable land are useful to pollinators depends in part on the temporal pattern of flowering and insect flight activity. We compiled citizen science data on 54 bees and hoverflies typical of agricultural areas in southern Sweden, as well as 24 flowering weed species classified as pollinator-friendly in the sense that they provide nectar and/or pollen to pollinators. The flight periods of the bees and hoverflies varied greatly, but there were also some consistent differences between the four groups studied. The first group to fly were the early flying solitary bees (7 species), followed by the social bees (18 species). In contrast, other solitary bees (11 species) and hoverflies (22 species) flew later in the summer. Solitary bees had the shortest flight periods, while social bees and hoverflies had longer flight periods. Flowering of weed species also varied greatly between species, with weeds classified as winter annuals (e.g., germinating in autumn) starting early together with germination generalists (species that can germinate in both autumn and spring). Summer annuals (spring germinators) and perennials started flowering about a month later. Germination generalists had a much longer flowering period than the others. Weekly pollinator records were in most cases significantly explained by weed records. Apart from early flying solitary bees, all models showed strong positive relationships. The overall best explanatory variable was the total number of weeds, with a weight assigned to each species based on its potential as a nectar/pollen source. This suggests that agricultural weeds in Sweden provide a continuous potential supply of nectar and pollen throughout the flight season of most pollinators.

Genetic structure, UV-vision, wing coloration and size coincide with colour polymorphism in Fabriciana adippe butterflies.

Colour polymorphisms have long served as model systems in evolutionary studies and continue to inform about processes involved in the origin and dynamics of biodiversity. Modern sequencing tools allow for evaluating whether phenotypic differences between morphs reflect genetic differentiation rather than developmental plasticity, and for investigating whether polymorphisms represent intermediate stages of diversification towards speciation. We investigated phenotypic and genetic differentiation between two colour morphs of the butterfly Fabriciana adippe using a combination of ddRAD-sequencing and comparisons of body size, colour patterns and optical properties of bright wing spots. The silvery-spotted adippe form had larger and darker wings and reflected UV light, while the yellow cleodoxa form displayed more green scales and reflected very little UV, showcasing that they constitute distinct and alternative integrated phenotypes. Genomic analyses revealed genetic structuring according to source population, and to colour morph, suggesting that the phenotypic differentiation reflects evolutionary modifications. We report 17 outlier loci associated with colour morph, including ultraviolet-sensitive visual pigment (UVRh1), which is associated with intraspecific communication and mate choice in butterflies. Together with the demonstration that the wings of the adippe (but essentially not the cleodoxa) morph reflect UV light, that UV reflectance is higher in females than males and that morphs differ in wing size, this suggests that these colour morphs might represent genetically integrated phenotypes, possibly adapted to different microhabitats. We propose that non-random mating might contribute to the differentiation and maintenance of the polymorphism.

Anthropogenic Influence on Moth Populations: A Comparative Study in Southern Sweden.

As moths are vital components of ecosystems and serve as important bioindicators, understanding the dynamics of their communities and the factors influencing these dynamics, such as anthropogenic impacts, is crucial to understand the ecological processes. Our study focuses on two provinces in southern Sweden, Västergötland and Småland, where we used province records from 1974 to 2019 in combination with light traps (in 2020) to record the presence and abundance of moth species, subsequently assessing species traits to determine potential associations with their presence in anthropogenically modified landscapes. This study design provides a unique opportunity to assess temporal changes in moth communities and their responses to shifts in environmental conditions, including anthropogenic impacts. Across the Västergötland and Småland provinces in Sweden, we recorded 776 moth taxa belonging to fourteen different taxonomic families of mainly Macroheterocera. We captured 44% and 28% of the total moth species known from these provinces in our traps in Borås (Västergötland) and Kalmar (Småland), respectively. In 2020, the species richness and abundance were higher in Borås than in Kalmar, while the Shannon and Simpson diversity indices revealed a higher species diversity in Kalmar. Between 1974 and 2019, the colonisation rates of the provinces increased faster in Småland. Ninety-three species were found to have colonised these provinces since 1974, showing that species richness increased over the study period. We reveal significant associations between the probability of a species being present in the traps and distinct traits compared to a provincial species pool. Traits over-represented in the traps included species with a high variation in colour patterns, generalist habitat preferences, extended flight periods, lower host plant specificity, and overwintering primarily as eggs. Our findings underscore the ongoing ecological filtering that favours certain species-specific traits. This study sheds light on the roles of climate change and anthropogenic impacts in shaping moth biodiversity, offers key insights into the ecological processes involved, and can guide future conservation efforts.

Century-long butterfly range expansions in northern Europe depend on climate, land use and species traits.

Climate change is an important driver of range shifts and community composition changes. Still, little is known about how the responses are influenced by the combination of land use, species interactions and species traits. We integrate climate and distributional data for 131 butterfly species in Sweden and Finland and show that cumulative species richness has increased with increasing temperature over the past 120 years. Average provincial species richness increased by 64% (range 15-229%), from 46 to 70. The rate and direction of range expansions have not matched the temperature changes, in part because colonisations have been modified by other climatic variables, land use and vary according to species characteristics representing ecological generalisation and species interactions. Results emphasise the role of a broad ecological filtering, whereby a mismatch between environmental conditions and species preferences limit the ability to disperse and establish populations in emerging climates and novel areas, with potentially widespread implications for ecosystem functioning.

An Approach to the Impact Simulation on Foamed Injection Molded Polypropylene Parts: An Example of Application in the Automotive Industry.

An approach to the simulation of foamed injection molded Polypropylene parts subjected to impact loading is presented in this paper. The proposed method, which considers strain-rate-dependent material properties and the possible occurrence of fracture, is, in particular, suitable for parts manufactured with core-back technology. The method was developed to be used within the functionality of a commercial Finite Element solver using a shell-type element mesh. The material model is based on a three-layer structure, with two compact skin layers and a foamed core layer made of expanded material. The properties of the foamed material are assumed as those of the compact grade scaled by a suitable factor, which is identified via inverse engineering on a set of bending tests executed on specimens having different foam densities. The fracture of the material is then predicted using a damage model which considers the effects of triaxiality. The approach is then validated on industrial parts from the automotive sector, subjected to impact in a component test. Despite the simplicity of the presented approach, which makes this method suitable for industrial applications and especially for early-stage design, the validation shows a sufficiently accurate simulation of part behavior under the impact, with a reasonable prediction of damage and fracture.

Associations of 16-Year Population Dynamics in Range-Expanding Moths with Temperature and Years since Establishment.

Parallel to the widespread decline of plants and animals, there is also an ongoing expansion of many species, which is especially pronounced in certain taxonomic groups and in northern latitudes. In order to inform an improved understanding of population dynamics in range-expanding taxa, we studied species richness, abundance and population growth in a sample of 25,138 individuals representing 107 range-expanding moth species at three light-trap sites in southeastern Sweden over 16 years (from 2005 to 2020) in relation to temperature and years since colonisation. Species richness and average abundance across range-expanding moths increased significantly over time, indicating a continuous influx of species expanding their ranges northward. Furthermore, average abundance and population growth increased significantly with increasing average ambient air temperature during the recording year, and average abundance also increased significantly with increasing temperature during the previous year. In general, population growth increased between years (growth rate > 1), although the population growth rate decreased significantly in association with years since colonisation. These findings highlight that, in contrast to several other studies in different parts of the world, species richness and abundance have increased in southeastern Sweden, partly because the warming climate enables range-expanding moths to realise their capacity for rapid distribution shifts and population growth. This may lead to fast and dramatic changes in community composition, with consequences for species interactions and the functioning of ecosystems. These findings are also of applied relevance for agriculture and forestry in that they can help to forecast the impacts of future invasive pest species.

Linking large-scale genetic structure of three Argynnini butterfly species to geography and environment.

Understanding which factors and processes are associated with genetic differentiation within and among species remains a major goal in evolutionary biology. To explore differences and similarities in genetic structure and its association with geographical and climatic factors in sympatric sister species, we conducted a large-scale (>32° latitude and >36° longitude) comparative phylogeographical study on three Argynnini butterfly species (Speyeria aglaja, Fabriciana adippe and F. niobe) that have similar life histories, but differ in ecological generalism and dispersal abilities. Analyses of nuclear (ddRAD-sequencing derived SNP markers) and mitochondrial (COI sequences) data revealed differences between species in genetic structure and how genetic differentiation was associated with climatic factors (temperature, solar radiation, precipitation, wind speed). Geographical proximity accounted for much of the variation in nuclear and mitochondrial structure and evolutionary relationships in F. adippe and F. niobe, but only explained the pattern observed in the nuclear data in S. aglaja, for which mitonuclear discordance was documented. In all species, Iberian and Balkan individuals formed genetic clusters, suggesting isolation in glacial refugia and limited postglacial expansion. Solar radiation and precipitation were associated with the genetic structure on a regional scale in all species, but the specific combinations of environmental and geographical factors linked to variation within species were unique, pointing to species-specific responses to common environments. Our findings show that the species share similar colonization histories, and that the same ecological factors, such as niche breadth and dispersal capacity, covary with genetic differentiation within these species to some extent, thereby highlighting the importance of comparative phylogeographical studies in sympatric sister species.

Differences in phenology, daily timing of activity, and associations of temperature utilization with survival in three threatened butterflies.

We used observational data collected during a mark-recapture study that generated a total of 7503 captures of 6108 unique individuals representing three endangered butterfly species to quantify inter-and intraindividual variation in temperature utilization and examine how activity patterns vary according to season, time of day, and ambient temperature. The Marsh Fritillary, the Apollo, and the Large Blue differed in utilized temperatures and phenology. Their daily activity patterns responded differently to temperature, in part depending on whether they were active in the beginning, middle or end of the season, in part reflecting interindividual variation and intraindividual flexibility, and in part owing to differences in ecology, morphology, and colouration. Activity temperatures varied over the season, and the Apollo and the Large Blue were primarily active at the highest available ambient temperatures (on the warmest days and during the warmest part of the day). The Marsh Fritillary was active early in the season and decreased activity during the highest temperatures. The relationship between individual lifespan and the average temperature was qualitatively different in the three species pointing to species-specific selection. Lifespan increased with an increasing range of utilized temperatures in all species, possibly reflecting that intra-individual flexibility comes with a general survival benefit.

Urban moth communities suggest that life in the city favours thermophilic multi-dimensional generalists.

Biodiversity is challenged worldwide by exploitation, global warming, changes in land use and increasing urbanization. It is hypothesized that communities in urban areas should consist primarily of generalist species with broad niches that are able to cope with novel, variable, fragmented, warmer and unpredictable environments shaped by human pressures. We surveyed moth communities in three cities in northern Europe and compared them with neighbouring moth assemblages constituting species pools of potential colonizers. We found that urban moth communities consisted of multi-dimensional generalist species that had larger distribution ranges, more variable colour patterns, longer reproductive seasons, broader diets, were more likely to overwinter as an egg, more thermophilic, and occupied more habitat types compared with moth communities in surrounding areas. When body size was analysed separately, results indicated that city occupancy was associated with larger size, but this effect disappeared when body size was analysed together with the other traits. Our findings indicate that urbanization imposes a spatial filtering process in favour of thermophilic species characterized by high intraspecific diversity and multi-dimensional generalist lifestyles over specialized species with narrow niches.

Declines in moth populations stress the need for conserving dark nights.

Given the global continuous rise, artificial light at night is often considered a driving force behind moth population declines. Although negative effects on individuals have been shown, there is no evidence for effects on population sizes to date. Therefore, we compared population trends of Dutch macromoth fauna over the period 1985-2015 between moth species that differ in phototaxis and adult circadian rhythm. We found that moth species that show positive phototaxis or are nocturnally active have stronger negative population trends than species that are not attracted to light or are diurnal species. Our results indicate that artificial light at night is an important factor in explaining declines in moth populations in regions with high artificial night sky brightness. Our study supports efforts to reduce the impacts of artificial light at night by promoting lamps that do not attract insects and reduce overall levels of illumination in rural areas to reverse declines of moth populations.

Faster poleward range shifts in moths with more variable colour patterns.

Range shifts have been documented in many organisms, and climate change has been implicated as a contributing driver of latitudinal and altitudinal range modifications. However, little is known about what species trait(s) allow for faster environmental tracking and improved capacity for distribution expansions. We used data for 416 species of moths, and show that range limits in Sweden have shifted to the north by on average 52.4 km per decade between 1973 and 2014. When also including non-expanding species, average expansion rate was 23.2 km per decade. The rate of boundary shifts increased with increasing levels of inter-individual variation in colour patterns and decreased with increasing latitude. The association with colour patterns indicate that variation in this functionally important trait enables species to cope with novel and changing conditions. Northern range limits also increased with average abundance and decreased with increasing year-to-year abundance fluctuations, implicating production of dispersers as a driver of range dynamics. Studies of terrestrial animals show that rates of poleward shifts differ between taxonomic groups, increase over time, and depend on study duration and latitude. Knowledge of how distribution shifts change with time, location, and species characteristics may improve projections of responses to climate change and aid the protection of biodiversity.

Predicting bee community responses to land-use changes: Effects of geographic and taxonomic biases.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

Functional traits help to explain half-century long shifts in pollinator distributions.

Changes in climate and land use can have important impacts on biodiversity. Species respond to such environmental modifications by adapting to new conditions or by shifting their geographic distributions towards more suitable areas. The latter might be constrained by species' functional traits that influence their ability to move, reproduce or establish. Here, we show that functional traits related to dispersal, reproduction, habitat use and diet have influenced how three pollinator groups (bees, butterflies and hoverflies) responded to changes in climate and land-use in the Netherlands since 1950. Across the three pollinator groups, we found pronounced areal range expansions (>53%) and modelled range shifts towards the north (all taxa: 17-22 km), west (bees: 14 km) and east (butterflies: 11 km). The importance of specific functional traits for explaining distributional changes varied among pollinator groups. Larval diet preferences (i.e. carnivorous vs. herbivorous/detritivorous and nitrogen values of host plants, respectively) were important for hoverflies and butterflies, adult body size for hoverflies, and flight period length for all groups. Moreover, interactions among multiple traits were important to explain species' geographic range shifts, suggesting that taxon-specific multi-trait analyses are needed to predict how global change will affect biodiversity and ecosystem services.

Variable coloration is associated with dampened population fluctuations in noctuid moths.

Theory and recent reviews state that greater genetic and phenotypic variation should be beneficial for population abundance and stability. Experimental evaluations of this prediction are rare, of short duration and conducted under controlled environmental settings. The question whether greater diversity in functionally important traits stabilizes populations under more complex ecological conditions in the wild has not been systematically evaluated. Moths are mainly nocturnal, with a large variation in colour patterns among species, and constitute an important food source for many types of organisms. Here, we report the results of a long-term (2003-2013) monitoring study of 115 100 noctuid moths from 246 species. Analysis of time-series data provide rare evidence that species with higher levels of inter-individual variation in colour pattern have higher average abundances and undergo smaller between-year fluctuations compared with species having less variable colour patterns. The signature of interspecific temporal synchronization of abundance fluctuations was weak, suggesting that the dynamics were driven by species-specific biotic interactions rather than by some common, density-independent factor(s). We conclude that individual variation in colour patterns dampens population abundance fluctuations, and suggest that this may partly reflect that colour pattern polymorphism provides protection from visually oriented predators and parasitoids.

Surveying moths using light traps: effects of weather and time of year.

Light trapping is an ideal method for surveying nocturnal moths, but in the absence of standardised survey methods effects of confounding factors may impede interpretation of the acquired data. We explored the influence of weather, time of year, and light source on nightly catches of macro moths in light traps, and compared four strategies for sampling by estimating observed species richness using rarefaction. We operated two traps with different light sources for 225 consecutive nights from mid-March to the end of October in eastern Germany in 2011. In total, 49 472 individuals of 372 species were recorded. Species richness and abundance per night were mainly influenced by night temperature, humidity and lamp type. With a limited sample size (<10 nights) it was slightly better to concentrate sampling on the warmest summer nights, but with more sampling nights it was slightly better to sample during the warmest nights in each month (March to October). By exploiting the higher moth activity during warm nights and an understanding of the species' phenology, it is possible to increase the number of species caught and reduce effects of confounding abiotic factors.

Population fluctuations and synchrony of grassland butterflies in relation to species traits.

Population fluctuations and synchrony influence population persistence; species with larger fluctuations and more synchronised population fluctuations face higher extinction risks. Here, we analyse the effect of diet specialisation, mobility, length of the flight period, and distance to the northern edge of the species' distribution in relation to between-year population fluctuations and synchrony of butterfly species. All butterfly species associated with grasslands were surveyed over five successive years at 19 grassland sites in a forest-dominated landscape (50 km(2)) in southern Sweden. At both the local and regional level, we found larger population fluctuations in species with longer flight periods. Population fluctuations were more synchronous among localities in diet specialists. Species with a long flight period might move more to track nectar resources compared to species with shorter flight period, and if nectar sources vary widely between years and localities it may explain that population fluctuations increase with increasing flight length. Diet generalists can use different resources (in this case host plants) at different localities and this can explain the lower synchrony in population fluctuations among generalist species. Higher degree of synchrony is one possible explanation for the higher extinction risks that have been observed for more specialised species. Therefore, diet specialists are more often threatened and require more conservation efforts than generalists.

High population variability and source-sink dynamics in a solitary bee species.

Although solitary bees are considered to play key roles in ecosystem functions, surprisingly few studies have explored their population dynamics. We investigated the population dynamics of a rare, declining, solitary bee (Andrena humilis) in a landscape of 80 km2 in southern Sweden from 2003 to 2011. Only one population was persistent throughout all years studied; most likely this population supplied the surrounding landscape with 11 smaller, temporary local populations. Despite stable pollen availability, the size of the persistent population fluctuated dramatically in a two-year cycle over the nine years, with 490-1230 nests in odd-numbered years and 21-48 nests in even-numbered years. These fluctuations were not significantly related to climatic variables or pollen availability. Nineteen colonization and 14 extinction events were recorded. Occupancy decreased with distance from the persistent population and increased with increasing resource (pollen) availability. There were significant positive correlations between the size of the persistent population and patch occupancy and colonization. Colonizations were generally more common in patches closer to the persistent population, whereas extinctions were independent of distance from the persistent population. Our results highlight the complex population dynamics that exist for this solitary bee species, which could be due to source-sink dynamics, a prolonged diapause, or can represent a bet-hedging strategy to avoid natural enemies and survive in small habitat patches. If large fluctuations in solitary bee populations prove to be widespread, it will have important implications for interpreting ecological relationships, bee conservation, and pollination.

With that diet, you will go far: trait-based analysis reveals a link between rapid range expansion and a nitrogen-favoured diet.

Recent global change has had a substantial influence on the distribution of organisms, and many species are currently expanding their ranges. To evaluate the underlying processes, long-term data with good geographic resolution are essential. One important but generally overlooked data source is offered by the taxon-specific national catalogues of first provincial records that are kept in many countries. Here, we use such data to quantify trait-based influences on range expansion in Swedish butterflies and moths between 1973 and 2010. Of 282 species meeting pre-defined quality criteria, 170 expanded their northern range margin, with a mean expansion rate of 2.7 km per year. The analyses demonstrate that habitat and diet generalists, forest species and species active during warm conditions have expanded their ranges more rapidly than other species. Notably, range expansion in diet specialists was positively related to a nitrogen-favoured larval diet, an effect not found among oligo- or polyphagous species. In contrast to the general view, this shows that specialist species can undergo rapid range expansion. We suggest that increased areas of nitrogen-rich habitat, and increased availability of a nitrogen-favoured diet, are among the most important drivers of range expansions, potentially having far-reaching consequences for a wide variety of organisms.

Species-area relationships are controlled by species traits.

The species-area relationship (SAR) is one of the most thoroughly investigated empirical relationships in ecology. Two theories have been proposed to explain SARs: classical island biogeography theory and niche theory. Classical island biogeography theory considers the processes of persistence, extinction, and colonization, whereas niche theory focuses on species requirements, such as habitat and resource use. Recent studies have called for the unification of these two theories to better explain the underlying mechanisms that generates SARs. In this context, species traits that can be related to each theory seem promising. Here we analyzed the SARs of butterfly and moth assemblages on islands differing in size and isolation. We tested whether species traits modify the SAR and the response to isolation. In addition to the expected overall effects on the area, traits related to each of the two theories increased the model fit, from 69% up to 90%. Steeper slopes have been shown to have a particularly higher sensitivity to area, which was indicated by species with restricted range (slope = 0.82), narrow dietary niche (slope= 0.59), low abundance (slope= 0.52), and low reproductive potential (slope = 0.51). We concluded that considering species traits by analyzing SARs yields considerable potential for unifying island biogeography theory and niche theory, and that the systematic and predictable effects observed when considering traits can help to guide conservation and management actions.

Both population size and patch quality affect local extinctions and colonizations.

Currently, the habitat of many species is fragmented, resulting in small local populations with individuals occasionally dispersing between the remaining habitat patches. In a solitary bee metapopulation, extinction probability was related to both local bee population sizes and pollen resources measured as host plant population size. Patch size, on the other hand, had no additional predictive power. The turnover rate of local bee populations in 63 habitat patches over 4 years was high, with 72 extinction events and 31 colonization events, but the pollen plant population was stable with no extinctions or colonizations. Both pollen resources and bee populations had strong and independent effects on extinction probability, but connectivity was not of importance. Colonizations occurred more frequently within larger host plant populations. For metapopulation survival of the bee, large pollen plant populations are essential, independent of current bee population size.