Recent range shifts of moths, butterflies, and birds are driven by the breadth of their climatic niche.

Species are altering their ranges as a response to climate change, but the magnitude and direction of observed range shifts vary considerably among species. The ability to persist in current areas and colonize new areas plays a crucial role in determining which species will thrive and which decline as climate change progresses. Several studies have sought to identify characteristics, such as morphological and life-history traits, that could explain differences in the capability of species to shift their ranges together with a changing climate. These characteristics have explained variation in range shifts only sporadically, thus offering an uncertain tool for discerning responses among species. As long-term selection to past climates have shaped species' tolerances, metrics describing species' contemporary climatic niches may provide an alternative means for understanding responses to on-going climate change. Species that occur in a broader range of climatic conditions may hold greater tolerance to climatic variability and could therefore more readily maintain their historical ranges, while species with more narrow tolerances may only persist if they are able to shift in space to track their climatic niche. Here, we provide a first-filter test of the effect of climatic niche dimensions on shifts in the leading range edges in three relatively well-dispersing species groups. Based on the realized changes in the northern range edges of 383 moth, butterfly, and bird species across a boreal 1,100 km latitudinal gradient over c. 20 years, we show that while most morphological or life-history traits were not strongly connected with range shifts, moths and birds occupying a narrower thermal niche and butterflies occupying a broader moisture niche across their European distribution show stronger shifts towards the north. Our results indicate that the climatic niche may be important for predicting responses under climate change and as such warrants further investigation of potential mechanistic underpinnings.

Mechanisms underpinning community stability along a latitudinal gradient: Insights from a niche-based approach.

At large scales, the mechanisms underpinning stability in natural communities may vary in importance due to changes in species composition, mean abundance, and species richness. Here we link species characteristics (niche positions) and community characteristics (richness and abundance) to evaluate the importance of stability mechanisms in 156 butterfly communities monitored across three European countries and spanning five bioclimatic regions. We construct niche-based hierarchical structural Bayesian models to explain first differences in abundance, population stability, and species richness between the countries, and then explore how these factors impact community stability both directly and indirectly (via synchrony and population stability). Species richness was partially explained by the position of a site relative to the niches of the species pool, and species near the centre of their niche had higher average population stability. The differences in mean abundance, population stability, and species richness then influenced how much variation in community stability they explained across the countries. We found, using variance partitioning, that community stability in Finnish communities was most influenced by community abundance, whereas this aspect was unimportant in Spain with species synchrony explaining most variation; the UK was somewhat intermediate with both factors explaining variation. Across all countries, the diversity-stability relationship was indirect with species richness reducing synchrony which increased community stability, with no direct effects of species richness. Our results suggest that in natural communities, biogeographical variation observed in key drivers of stability, such as population abundance and species richness, leads to community stability being limited by different factors and that this can partially be explained due to the niche characteristics of the European butterfly assemblage.

Local adaptation to climate anomalies relates to species phylogeny.

Climatic anomalies are increasing in intensity and frequency due to rapid rates of global change, leading to increased extinction risk for many species. The impacts of anomalies are likely to vary between species due to different degrees of sensitivity and extents of local adaptation. Here, we used long-term butterfly monitoring data of 143 species across six European bioclimatic regions to show how species' population dynamics have responded to local or globally-calculated climatic anomalies, and how species attributes mediate these responses. Contrary to expectations, degree of apparent local adaptation, estimated from the relative population sensitivity to local versus global anomalies, showed no associations with species mobility or reproductive rate but did contain a strong phylogenetic signal. The existence of phylogenetically-patterned local adaptation to climate has important implications for forecasting species responses to current and future climatic conditions and for developing appropriate conservation practices.

Urbanization extends flight phenology and leads to local adaptation of seasonal plasticity in Lepidoptera.

Urbanization is gaining force globally, which challenges biodiversity, and it has recently also emerged as an agent of evolutionary change. Seasonal phenology and life cycle regulation are essential processes that urbanization is likely to alter through both the urban heat island effect (UHI) and artificial light at night (ALAN). However, how UHI and ALAN affect the evolution of seasonal adaptations has received little attention. Here, we test for the urban evolution of seasonal life-history plasticity, specifically changes in the photoperiodic induction of diapause in two lepidopterans, Pieris napi (Pieridae) and Chiasmia clathrata (Geometridae). We used long-term data from standardized monitoring and citizen science observation schemes to compare yearly phenological flight curves in six cities in Finland and Sweden to those of adjacent rural populations. This analysis showed for both species that flight seasons are longer and end later in most cities, suggesting a difference in the timing of diapause induction. Then, we used common garden experiments to test whether the evolution of the photoperiodic reaction norm for diapause could explain these phenological changes for a subset of these cities. These experiments demonstrated a genetic shift for both species in urban areas toward a lower daylength threshold for direct development, consistent with predictions based on the UHI but not ALAN. The correspondence of this genetic change to the results of our larger-scale observational analysis of in situ flight phenology indicates that it may be widespread. These findings suggest that seasonal life cycle regulation evolves in urban ectotherms and may contribute to ecoevolutionary dynamics in cities.

Combining range and phenology shifts offers a winning strategy for boreal Lepidoptera.

Species can adapt to climate change by adjusting in situ or by dispersing to new areas, and these strategies may complement or enhance each other. Here, we investigate temporal shifts in phenology and spatial shifts in northern range boundaries for 289 Lepidoptera species by using long-term data sampled over two decades. While 40% of the species neither advanced phenology nor moved northward, nearly half (45%) used one of the two strategies. The strongest positive population trends were observed for the minority of species (15%) that both advanced flight phenology and shifted their northern range boundaries northward. We show that, for boreal Lepidoptera, a combination of phenology and range shifts is the most viable strategy under a changing climate. Effectively, this may divide species into winners and losers based on their propensity to capitalize on this combination, with potentially large consequences on future community composition.

Habitat amount and distribution modify community dynamics under climate change.

Habitat fragmentation may present a major impediment to species range shifts caused by climate change, but how it affects local community dynamics in a changing climate has so far not been adequately investigated empirically. Using long-term monitoring data of butterfly assemblages, we tested the effects of the amount and distribution of semi-natural habitat (SNH), moderated by species traits, on climate-driven species turnover. We found that spatially dispersed SNH favoured the colonisation of warm-adapted and mobile species. In contrast, extinction risk of cold-adapted species increased in dispersed (as opposed to aggregated) habitats and when the amount of SNH was low. Strengthening habitat networks by maintaining or creating stepping-stone patches could thus allow warm-adapted species to expand their range, while increasing the area of natural habitat and its spatial cohesion may be important to aid the local persistence of species threatened by a warming climate.

Landscape simplification weakens the association between terrestrial producer and consumer diversity in Europe.

Land-use change is one of the primary drivers of species loss, yet little is known about its effect on other components of biodiversity that may be at risk. Here, we ask whether, and to what extent, landscape simplification, measured as the percentage of arable land in the landscape, disrupts the functional and phylogenetic association between primary producers and consumers. Across seven European regions, we inferred the potential associations (functional and phylogenetic) between host plants and butterflies in 561 seminatural grasslands. Local plant diversity showed a strong bottom-up effect on butterfly diversity in the most complex landscapes, but this effect disappeared in simple landscapes. The functional associations between plant and butterflies are, therefore, the results of processes that act not only locally but are also dependent on the surrounding landscape context. Similarly, landscape simplification reduced the phylogenetic congruence among host plants and butterflies indicating that closely related butterflies become more generalist in the resources used. These processes occurred without any detectable change in species richness of plants or butterflies along the gradient of arable land. The structural properties of ecosystems are experiencing substantial erosion, with potentially pervasive effects on ecosystem functions and future evolutionary trajectories. Loss of interacting species might trigger cascading extinction events and reduce the stability of trophic interactions, as well as influence the longer term resilience of ecosystem functions. This underscores a growing realization that species richness is a crude and insensitive metric and that both functional and phylogenetic associations, measured across multiple trophic levels, are likely to provide additional and deeper insights into the resilience of ecosystems and the functions they provide.

Weather explains high annual variation in butterfly dispersal.

Weather conditions fundamentally affect the activity of short-lived insects. Annual variation in weather is therefore likely to be an important determinant of their between-year variation in dispersal, but conclusive empirical studies are lacking. We studied whether the annual variation of dispersal can be explained by the flight season's weather conditions in a Clouded Apollo (Parnassius mnemosyne) metapopulation. This metapopulation was monitored using the mark-release-recapture method for 12 years. Dispersal was quantified for each monitoring year using three complementary measures: emigration rate (fraction of individuals moving between habitat patches), average residence time in the natal patch, and average distance moved. There was much variation both in dispersal and average weather conditions among the years. Weather variables significantly affected the three measures of dispersal and together with adjusting variables explained 79-91% of the variation observed in dispersal. Different weather variables became selected in the models explaining variation in three dispersal measures apparently because of the notable intercorrelations. In general, dispersal rate increased with increasing temperature, solar radiation, proportion of especially warm days, and butterfly density, and decreased with increasing cloudiness, rainfall, and wind speed. These results help to understand and model annually varying dispersal dynamics of species affected by global warming.

Impacts of land cover data selection and trait parameterisation on dynamic modelling of species' range expansion.

Dynamic models for range expansion provide a promising tool for assessing species' capacity to respond to climate change by shifting their ranges to new areas. However, these models include a number of uncertainties which may affect how successfully they can be applied to climate change oriented conservation planning. We used RangeShifter, a novel dynamic and individual-based modelling platform, to study two potential sources of such uncertainties: the selection of land cover data and the parameterization of key life-history traits. As an example, we modelled the range expansion dynamics of two butterfly species, one habitat specialist (Maniola jurtina) and one generalist (Issoria lathonia). Our results show that projections of total population size, number of occupied grid cells and the mean maximal latitudinal range shift were all clearly dependent on the choice made between using CORINE land cover data vs. using more detailed grassland data from three alternative national databases. Range expansion was also sensitive to the parameterization of the four considered life-history traits (magnitude and probability of long-distance dispersal events, population growth rate and carrying capacity), with carrying capacity and magnitude of long-distance dispersal showing the strongest effect. Our results highlight the sensitivity of dynamic species population models to the selection of existing land cover data and to uncertainty in the model parameters and indicate that these need to be carefully evaluated before the models are applied to conservation planning.

Density of insect-pollinated grassland plants decreases with increasing surrounding land-use intensity.

Pollinator declines have raised concerns about the persistence of plant species that depend on insect pollination, in particular by bees, for their reproduction. The impact of pollinator declines remains unknown for species-rich plant communities found in temperate seminatural grasslands. We investigated effects of land-use intensity in the surrounding landscape on the distribution of plant traits related to insect pollination in 239 European seminatural grasslands. Increasing arable land use in the surrounding landscape consistently reduced the density of plants depending on bee and insect pollination. Similarly, the relative abundance of bee-pollination-dependent plants increased with higher proportions of non-arable agricultural land (e.g. permanent grassland). This was paralleled by an overall increase in bee abundance and diversity. By isolating the impact of the surrounding landscape from effects of local habitat quality, we show for the first time that grassland plants dependent on insect pollination are particularly susceptible to increasing land-use intensity in the landscape.

Higher mobility of butterflies than moths connected to habitat suitability and body size in a release experiment.

Mobility is a key factor determining lepidopteran species responses to environmental change. However, direct multispecies comparisons of mobility are rare and empirical comparisons between butterflies and moths have not been previously conducted. Here, we compared mobility between butterflies and diurnal moths and studied species traits affecting butterfly mobility. We experimentally marked and released 2011 butterfly and 2367 moth individuals belonging to 32 and 28 species, respectively, in a 25 m × 25 m release area within an 11-ha, 8-year-old set-aside field. Distance moved and emigration rate from the release habitat were recorded by species. The release experiment produced directly comparable mobility data in 18 butterfly and 9 moth species with almost 500 individuals recaptured. Butterflies were found more mobile than geometroid moths in terms of both distance moved (mean 315 m vs. 63 m, respectively) and emigration rate (mean 54% vs. 17%, respectively). Release habitat suitability had a strong effect on emigration rate and distance moved, because butterflies tended to leave the set-aside, if it was not suitable for breeding. In addition, emigration rate and distance moved increased significantly with increasing body size. When phylogenetic relatedness among species was included in the analyses, the significant effect of body size disappeared, but habitat suitability remained significant for distance moved. The higher mobility of butterflies than geometroid moths can largely be explained by morphological differences, as butterflies are more robust fliers. The important role of release habitat suitability in butterfly mobility was expected, but seems not to have been empirically documented before. The observed positive correlation between butterfly size and mobility is in agreement with our previous findings on butterfly colonization speed in a long-term set-aside experiment and recent meta-analyses on butterfly mobility.

Model for quantifying the synergies between farmland biodiversity conservation and water protection at catchment scale.

This paper studies joint provision of two environmental non-market commodities related to agriculture: biodiversity conservation and water protection. We provide an optimising tool for analysing spatial dependencies of multifunctional agriculture at catchment scale. We show that efficiency gains can be achieved by spatial allocation and choice of the type of vegetation. In particular, inclusion of meadow nectar plants in the founding grass seed mixture of set-asides was found out to be an economically efficient measure to promote biodiversity and water protection on warm, steep slopes.

Environmental factors driving the effectiveness of European agri-environmental measures in mitigating pollinator loss--a meta-analysis.

In Europe, agri-environmental schemes (AES) have been introduced in response to concerns about farmland biodiversity declines. Yet, as AES have delivered variable results, a better understanding of what determines their success or failure is urgently needed. Focusing on pollinating insects, we quantitatively reviewed how environmental factors affect the effectiveness of AES. Our results suggest that the ecological contrast in floral resources created by schemes drives the response of pollinators to AES but that this response is moderated by landscape context and farmland type, with more positive responses in croplands (vs. grasslands) located in simple (vs. cleared or complex) landscapes. These findings inform us how to promote pollinators and associated pollination services in species-poor landscapes. They do not, however, present viable strategies to mitigate loss of threatened or endangered species. This indicates that the objectives and design of AES should distinguish more clearly between biodiversity conservation and delivery of ecosystem services.

Life-history traits predict species responses to habitat area and isolation: a cross-continental synthesis.

There is a lack of quantitative syntheses of fragmentation effects across species and biogeographic regions, especially with respect to species life-history traits. We used data from 24 independent studies of butterflies and moths from a wide range of habitats and landscapes in Europe and North America to test whether traits associated with dispersal capacity, niche breadth and reproductive rate modify the effect of habitat fragmentation on species richness. Overall, species richness increased with habitat patch area and connectivity. Life-history traits improved the explanatory power of the statistical models considerably and modified the butterfly species-area relationship. Species with low mobility, a narrow feeding niche and low reproduction were most strongly affected by habitat loss. This demonstrates the importance of considering life-history traits in fragmentation studies and implies that both species richness and composition change in a predictable manner with habitat loss and fragmentation.

Habitat fragmentation causes immediate and time-delayed biodiversity loss at different trophic levels.

Intensification or abandonment of agricultural land use has led to a severe decline of semi-natural habitats across Europe. This can cause immediate loss of species but also time-delayed extinctions, known as the extinction debt. In a pan-European study of 147 fragmented grassland remnants, we found differences in the extinction debt of species from different trophic levels. Present-day species richness of long-lived vascular plant specialists was better explained by past than current landscape patterns, indicating an extinction debt. In contrast, short-lived butterfly specialists showed no evidence for an extinction debt at a time scale of c. 40 years. Our results indicate that management strategies maintaining the status quo of fragmented habitats are insufficient, as time-delayed extinctions and associated co-extinctions will lead to further biodiversity loss in the future.

Extinction debt: a challenge for biodiversity conservation.

Local extinction of species can occur with a substantial delay following habitat loss or degradation. Accumulating evidence suggests that such extinction debts pose a significant but often unrecognized challenge for biodiversity conservation across a wide range of taxa and ecosystems. Species with long generation times and populations near their extinction threshold are most likely to have an extinction debt. However, as long as a species that is predicted to become extinct still persists, there is time for conservation measures such as habitat restoration and landscape management. Standardized long-term monitoring, more high-quality empirical studies on different taxa and ecosystems and further development of analytical methods will help to better quantify extinction debt and protect biodiversity.

Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands.

Metapopulation theory predicts that species richness and total population density of habitat specialists increase with increasing area and regional connectivity of the habitat. To test these predictions, we examined the relative contributions of habitat patch area, connectivity of the regional habitat network and local habitat quality to species richness and total density of butterflies and day-active moths inhabiting semi-natural grasslands. We studied butterflies and moths in 48 replicate landscapes situated in southwest Finland, including a focal patch and the surrounding network of other semi-natural grasslands within a radius of 1.5 km from the focal patch. By applying the method of hierarchical partitioning, which can distinguish between independent and joint contributions of individual explanatory variables, we observed that variables of the local habitat quality (e.g. mean vegetation height and nectar plant abundance) generally showed the highest independent effect on species richness and total density of butterflies and moths. Habitat area did not show a significant independent contribution to species richness and total density of butterflies and moths. The effect of habitat connectivity was observed only for total density of the declining butterflies and moths. These observations indicate that the local habitat quality is of foremost importance in explaining variation in species richness and total density of butterflies and moths. In addition, declining butterflies and moths have larger populations in well-connected networks of semi-natural grasslands. Our results suggest that, while it is crucial to maintain high-quality habitats by management, with limited resources it would be appropriate to concentrate grassland management and restoration to areas with well-connected grassland networks in which the declining species currently have their strongest populations.

An empirical test of a diffusion model: predicting clouded apollo movements in a novel environment.

Functional connectivity is a fundamental concept in conservation biology because it sets the level of migration and gene flow among local populations. However, functional connectivity is difficult to measure, largely because it is hard to acquire and analyze movement data from heterogeneous landscapes. Here we apply a Bayesian state-space framework to parameterize a diffusion-based movement model using capture-recapture data on the endangered clouded apollo butterfly. We test whether the model is able to disentangle the inherent movement behavior of the species from landscape structure and sampling artifacts, which is a necessity if the model is to be used to examine how movements depend on landscape structure. We show that this is the case by demonstrating that the model, parameterized with data from a reference landscape, correctly predicts movements in a structurally different landscape. In particular, the model helps to explain why a movement corridor that was constructed as a management measure failed to increase movement among local populations. We illustrate how the parameterized model can be used to derive biologically relevant measures of functional connectivity, thus linking movement data with models of spatial population dynamics.

New insights into butterfly-environment relationships using partitioning methods.

Variation partitioning and hierarchical partitioning are novel statistical approaches that provide deeper understanding of the importance of different explanatory variables for biodiversity patterns than traditional regression methods. Using these methods, the variation in occupancy and abundance of the clouded apollo butterfly (Parnassius mnemosyne L.) was decomposed into independent and joint effects of larval and adult food resources, microclimate and habitat quantity. The independent effect of habitat quantity variables (habitat area and connectivity) captured the largest fraction of the variation in the clouded apollo patterns, but habitat connectivity had a major contribution only for occupancy data. The independent effects of resources and microclimate were higher on butterfly abundance than on occupancy. However, a considerable amount of variation in the butterfly patterns was accounted for by the joint effects of predictors and may thus be causally related to two or all three groups of variables. Abundance of the butterfly in the surroundings of the focal grid cell had a significant effect in all analyses, independently of the effects of other predictors. Our results encourage wider applications of partitioning methods in biodiversity studies.

Use of forest corridors by boreal Xestia moths.

We studied movements of the noctuid moths Xestia speciosa and X. fennica in a fragmented forest landscape including old-growth spruce forest corridors in eastern Finland. We individually marked and released 1,796 adults of the habitat generalist X. speciosa and 683 individuals of X. fennica, which prefers old-growth spruce forests. Moths were recaptured with 60 sugar-bait traps that were placed within habitat corridors, in the non-habitat matrix and in surrounding old-growth forest patches. Both species favoured the corridors during their movements and tended to avoid entering the matrix. We analysed recapture rates and movements of the recaptured moths. Results suggest that X. speciosa moves longer distances than X. fennica. Though corridors clearly canalized movements by both species, the impact of old-growth forest corridors at the larger landscape scale was less clear in X. fennica. Movement behaviour of the two sexes differed more than that of the two species.