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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.

Species-rich semi-natural grasslands have been lost during the last century due to agricultural intensification. This has had large negative consequences for many specialised species, including grassland butterflies. To prevent further loss, management regimes in the remaining grasslands must maintain habitat quality over time, and we therefore need to understand the habitat preferences of specialised species and how different management regimes affect their survival. We studied the egg-laying preferences of the threatened marsh fritillary butterfly in relation to host plant properties, microclimate and management (grazing) on Gotland, Sweden. Moreover, we followed the survival of eggs and larvae from 27 egg batches during a period of 8 months (from June 2020 to March 2021) in grazed and ungrazed areas. We found 92 egg batches in total and the average number of eggs was 184.5. Egg-laying probability increased with increasing host plant size and abundance, and environmental variables associated with a warm microclimate (low grass cover, low vegetation height and south-facing edges). The 27 egg batches that were followed over time had on average 203 eggs in June. Roughly 28% of the eggs developed into larvae, and about 17% of these survived over the entire study period, resulting in an overall 4.7% survival. Egg survival was higher in ungrazed habitats compared with grazed; in March (post-hibernation), there were almost nine times more eggs in ungrazed habitats. This study highlights the complex habitat ecology of specialised butterflies and underscores the detrimental impact of intense grazing, advocating for rotational grazing or mowing regimes. We study oviposition preferences and survival of both eggs and larvae (post-hibernation) for the marsh fritillary, in relation to host plant availability, microclimate conditions and grazing management Egg-laying probability increased with increasing host plant size and abundance, and environmental variables associated with

1. Established butterfly survey methods that are useful for monitoring species that are widely distributed and numerous may be less accurate for more rare species. We therefore need new monitoring approaches.2. We describe a plot-based survey method, where butterflies or larvae nests are counted within 1-ha grid cells. The aim was to compare this grid method with more traditional transect counts and evaluate both methods in relation to high-quality capture-mark-release (CMR) population estimates (reflecting the 'true' population). We do this using data from a large population of the marsh fritillary butterfly in Sweden. Moreover, we followed the overall population trend from 2017 to 2021 for both adult butterflies and larvae.3. Results showed a higher detection probability using the grid method compared with transect counts, which for adult butterflies seem to be explained by time effort. Moreover, grid surveys of adult butterflies showed a clear significant relationship with the estimated 'true' population size from CMR, while transect counts did not. For larvae, both methods showed significant relationships with the estimated adult population size, but the grid method found 5.7 times more larvae. The overall popu-lation fluctuated significantly across years. In years with low densities, the transect method largely failed to detect the species.4. The grid method seems more reliable for detecting the marsh fritillary and for estimating its population size, and thus, tracking the population trend. We propose this novel method to be integrated into surveys and monitoring of biodiversity, especially when focusing on rare habitat specialists that are normally underrepre-sented in monitoring based on volunteer counts.

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.

Simple Summary There has been a widespread decline of many plants and animals driven at least partly by climate change. This pattern is not universal, and certain taxa are increasing in abundance and distribution. A better understanding of population dynamics and range expansions in different areas and how different taxa respond to changing temperatures is therefore important, as we are facing a warmer and more fluctuating climate in the future. In this study, we show that range-expanding moths in southeastern Sweden have increased their species richness over time and that abundance and population growth increase during years with higher temperatures. We also show that population growth in range-expanding moths is fastest in the first years after establishment in an area. These shifts in distribution and abundance of moths may lead to rapid and dramatic changes in community compositions, with potentially widespread consequences for species interactions and ecosystem functioning. 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.

Understanding how large-scale environmental variability may shape the distribution of phenotypic variation remains central to evolutionary biology. Across-species comparisons of trait variation alongside environmental gradients may offer valuable insights into how different species may respond to similar selective pressures. We conducted a comparative morphological study (>32° latitude and >47° longitude) on three closely related Argynnini butterfly species, Speyeria aglaja, Fabriciana adippe, and F. niobe. We measured wing size and coloration to assess (1) whether they respond similarly or differently to environmental factors (longitude, latitude, altitude, temperature, precipitation, solar radiation, wind speed); (2) if these factors correspond with those associated with the species’ genetic structure based on a previous study; and (3) whether correlations between phenotypic traits within individuals are species-specific. We found common and species-specific associations of climatic (precipitation, wind speed) and geographic (longitude, altitude) factors with the composite phenotypic variation. Wing size was associated with different variables in the studied species, while melanisation mainly increased in cooler regions in all species, suggesting that the need for temperature regulation is a strong selective pressure on melanisation. Wing size was associated with the genetic structure in all species, highlighting the functional importance of this trait. The environmental drivers associated with the phenotypic variation in S. aglaja and F. adippe were largely the same as those associated with their genetic structure, hinting at a genetic underpinning of the observed morphological variation due to local adaption. We report some distinct intraspecific trait correlations in S. aglaja and F. adippe, indicative of independent phenotypic integration. These phenotypes seem to be associated with protection against predators and thermal regulation in the respective ha

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.

Environmental and climatic changes are inducing population declines in numerous species. However, certain species demonstrate remarkable resilience, exhibiting both population growth and range expansion. This longitudinal study in Sweden carried out over two decades (2004–2023) examines the noctuid moths Mythimna albipuncta and Hoplodrina ambigua. Abundance and phenology data were gathered from three light traps in southeastern Sweden and integrated with distribution and phenology data from the Global Biodiversity Information Facility. In M. albipuncta, the distribution area expanded from 7 to 76 occupied grids (60 km2) and the abundance increased from 7 to 6136 individuals, while in H. ambigua, the distribution area expanded from 1 to 87 occupied grids and the abundance increased from 0 to 6937 individuals, during the course of the study. Furthermore, a positive yearly association was observed between the number of occupied grids and light trap abundance for each species. We also found significant extensions in the adult flight periods of more than 100 days in both species. Light traps emerged as an effective monitoring tool, with light trap abundance as a reliable proxy for distribution changes. Our findings demonstrate that the studied species cope very well with environmental and climatic changes. Given their role as dominant links between primary producers and higher trophic levels, abundance and distribution shifts of these ecological engineers have the potential to cascade up and down in the ecosystem.

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, V & auml;sterg & ouml;tland and Sm & aring;land, 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 V & auml;sterg & ouml;tland and Sm & aring;land 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 Bor & aring;s (V & auml;sterg & ouml;tland) and Kalmar (Sm & aring;land), respectively. In 2020, the species richness and abundance were higher in Bor & aring;s 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 Sm & aring;land. 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 specif

Understanding how large-scale environmental variability may shape the distribution of phenotypic variation remains central to evolutionary biology. Across-species comparisons of trait variation alongside environmental gradients may offer valuable insights into how different species may respond to similar selective pressures. We conducted a comparative morphological study (>32° latitude and >47° longitude) on three closely related Argynnini butterfly species, Speyeria aglaja, Fabriciana adippe, and F. niobe. We measured wing size and coloration to assess (1) whether they respond similarly or differently to environmental factors (longitude, latitude, altitude, temperature, precipitation, solar radiation, wind speed); (2) if these factors correspond with those associated with the species’ genetic structure based on a previous study; and (3) whether correlations between phenotypic traits within individuals are species-specific. We found common and species-specific associations of climatic (precipitation, wind speed) and geographic (longitude, altitude) factors with the composite phenotypic variation. Wing size was associated with different variables in the studied species, while melanisation mainly increased in cooler regions in all species, suggesting that the need for temperature regulation is a strong selective pressure on melanisation. Wing size was associated with the genetic structure in all species, highlighting the functional importance of this trait. The environmental drivers associated with the phenotypic variation in S. aglaja and F. adippe were largely the same as those associated with their genetic structure, hinting at a genetic underpinning of the observed morphological variation due to local adaption. We report some distinct intraspecific trait correlations in S. aglaja and F. adippe, indicative of independent phenotypic integration. These phenotypes seem to be associated with protection against predators and thermal regulation in the respective habitats

It remains unclear to what extent yearly weather variation and spatial variation in microclimate influences the outcome of interacting plant-animal species and whether responses differ between life stages. We collected data over several years on 46 ha on File Hajdar, Gotland, Sweden, and executed a complete mapping of larva nests (n = 776) and imago (n = 5,952) of the marsh fritillary butterfly Euphydryas aurinia and its host plant Succisa pratensis. The phenology of the butterflies and the major nectar plants visited varied among years. The duration of the adult flight period decreased with increasing ambient air temperatures. The density of butterflies, host plants, and host plant leaf size increased between years with increasing precipitation in the preceding year, and decreased with increasing average ambient air temperature in the preceding year. In 2021-2022 we deployed a unmanned aerial vehicle (UAV) with a high-resolution thermal sensor to measure spatial variation in surface temperatures in the study area. We found that survival from the egg to the larva stage increased with increasing surface temperature and host plant density. Host plants and larva nests generally occupied warmer microhabitats compared to imago butterflies. The results further suggested that the relationships linking surface temperature to the densities of imago, larva, host plants, and leaf size differed qualitatively between years. In 2017, larva nests and host plant density increased with increasing surface temperatures, and butterflies showed a non-linear response with a density peak at intermediate temperatures. As a result of the extreme drought in 2018 there was a reduction in maximum leaf size, and in the densities of plants, larvae, and butterflies. Moreover, the slopes of the relationships linking the density of larvae, butterflies, and plants to temperature shifted from linear positive to negative or curvilinear. Our findings demonstrate how yearly weather variation and heterog

In Europe, Natura 2000 sites should protect threatened target species and networks of habitats. The management of Natura 2000 grasslands is often financed by subsidized grazing as part of the Common Agricultural Policy (CAP). We studied the extent of CAP grazing for Natura 2000 management and how this affects a butterfly target species (the marsh fritillary) and floral resources. Based on extensive capture-mark-release studies from 2 years in >550 ha grid cells in a 225 km(2) landscape in Sweden that includes 15 Natura 2000 sites, we compared marsh fritillary occurrence probabilities and population densities in ungrazed and CAP-grazed habitats. Moreover, we analyzed how nectar resources and orchids were affected by CAP grazing based on plants records from 2347 sample plots. We estimated the proportion of butterfly habitats that were CAP-grazed within and outside Natura 2000 sites. In total, 10 453 and 4417 butterflies were marked in 2017 and 2019, respectively. The grid cell occurrence probability was 1.8 times higher and the population density was 2.3 times higher in ungrazed compared with CAP-grazed habitats in 2017, and the corresponding numbers for 2019 were 10 and 5.3 times higher, respectively. The number of flowering plants were on average 6.9 times higher and the density of orchids was 12.3 times higher in ungrazed habitats. Roughly, 30% (130 ha) of the marsh fritillary habitat was CAP grazed, and 97% of this grazing occurred within protected areas, of which 111 ha was situated within Natura 2000 area where the marsh fritillary is the target species. Alarmingly, we show that intense yearly CAP grazing, which is the dominant management strategy in all Natura 2000 sites, has devastating consequences for the target species and other aspects of biodiversity. Less intense management, which would benefit biodiversity, requires changes in the CAP, to allow more flexible payments for habitat management objectives and conservation of target species.

Extreme weather events are becoming more frequent due to climate change. We therefore need to understand how species respond to these events. In 2018, the worst drought ever recorded hit the island of Gotland in the Baltic Sea, which led to a major decline of the threatened marsh fritillary (Euphydryas aurinia) the succeeding summer. We compared yearly occupancy among 256 habitat patches between 2018 and 2020 and analysed the colonisation-extinction dynamics between 2019 and 2020 in relation to patch area, connectivity, and habitat quality. Moreover, using capture-mark-release data in two patch clusters differing in connectivity, we also compared population sizes before and after the drought and analysed 'population growth rates'. We also compared yearly abundance of host plants (Succisa pratensis) over time. Results show a remarkable recovery of the marsh fritillary. Both patch occupancy in 2020 and the number of individuals in the well-connected patch cluster were higher than before the drought. In contrast, host plants were fewer and smaller, which taken together suggest that the amount of food resources was roughly half in 2020 compared to the pre-drought conditions. Moreover, the butterfly population in the less connected patch cluster was eight times smaller compared to the population size before the drought. Local colonisations, extinctions, and population growth rates were explained by connectivity. The ability to quickly recover after extreme droughts is promising in times of climate change. The significance of connectivity for the population dynamics during recovery highlights the importance of maintaining well-connected patch networks.

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 degrees latitude and >36 degrees 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.