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.

Land sharing complements land sparing in the conservation of disturbance-dependent species.

Alteration of natural disturbances in human-modified landscapes has resulted in many disturbance-dependent species becoming rare. Conservation of such species requires efforts to maintain or recreate disturbance regimes. We compared benefits of confining efforts to habitats in protected areas (a form of land sparing) versus integrating them with general management of production land (a form of land sharing), using two examples: fire in forests and grazing in semi-natural grasslands. We reviewed empirical studies from the temperate northern hemisphere assessing effects of disturbances in protected and non-protected areas, and compiled information from organisations governing and implementing disturbances in Sweden. We found advantages with protection of areas related to temporal continuity and quality of disturbances, but the spatial extent of disturbances is higher on production land. This suggests that an approach where land sparing is complemented with land sharing will be most effective for preservation of disturbance-dependent species in forests and semi-natural grasslands.

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.

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.

From Agricultural Benefits to Aviation Safety: Realizing the Potential of Continent-Wide Radar Networks.

Migratory animals provide a multitude of services and disservices-with benefits or costs in the order of billions of dollars annually. Monitoring, quantifying, and forecasting migrations across continents could assist diverse stakeholders in utilizing migrant services, reducing disservices, or mitigating human-wildlife conflicts. Radars are powerful tools for such monitoring as they can assess directional intensities, such as migration traffic rates, and biomass transported. Currently, however, most radar applications are local or small scale and therefore substantially limited in their ability to address large-scale phenomena. As weather radars are organized into continent-wide networks and also detect "biological targets," they could routinely monitor aerial migrations over the relevant spatial scales and over the timescales required for detecting responses to environmental perturbations. To tap these unexploited resources, a concerted effort is needed among diverse fields of expertise and among stakeholders to recognize the value of the existing infrastructure and data beyond weather forecasting.

Climate regulation, energy provisioning and water purification: Quantifying ecosystem service delivery of bioenergy willow grown on riparian buffer zones using life cycle assessment.

Whilst life cycle assessment (LCA) boundaries are expanded to account for negative indirect consequences of bioenergy such as indirect land use change (ILUC), ecosystem services such as water purification sometimes delivered by perennial bioenergy crops are typically neglected in LCA studies. Consequential LCA was applied to evaluate the significance of nutrient interception and retention on the environmental balance of unfertilised energy willow planted on 50-m riparian buffer strips and drainage filtration zones in the Skåne region of Sweden. Excluding possible ILUC effects and considering oil heat substitution, strategically planted filter willow can achieve net global warming potential (GWP) and eutrophication potential (EP) savings of up to 11.9 Mg CO2e and 47 kg PO4e ha-1 year-1, respectively, compared with a GWP saving of 14.8 Mg CO2e ha-1 year-1 and an EP increase of 7 kg PO4e ha-1 year-1 for fertilised willow. Planting willow on appropriate buffer and filter zones throughout Skåne could avoid 626 Mg year-1 PO4e nutrient loading to waters.

Geographic and habitat origin influence biomass production and storage translocation in the clonal plant Aegopodium podagraria.

Through physiological integration, clonal plants can support ramets in unfavourable patches, exploit heterogeneously distributed resources and distribute resources that are taken up over large areas. Physiological integration generally increases in adverse conditions, but it is not well known which factors determine the evolution of physiological integration. The aim of this study was to investigate if clonal plants from Southern and Northern populations of the clonal herb Aegopodium podagraria differed in physiological integration in terms of translocation of carbon to the rhizomes, and in biomass production using a reciprocal transplant experiment. Aegopodium podagraria from shaded conditions have been suggested to share more resources than clones from open conditions and therefore, plants from forest and open populations within the Southern and Northern regions were included. The regional growing conditions greatly affected biomass production. Plants grown in North Sweden produced more biomass and allocated more biomass to shoots, while plants grown in South Sweden allocated more biomass to rhizomes. There was a regional origin effect as plants originating from North Sweden produced more biomass in both regions. Within the Northern region, plants from shaded habitats translocated more (14)C to the rhizomes, suggesting more storage there than in plants from open habitats. In addition to genetic differentiation in biomass production between Northern and Southern populations, probably as a response to a shorter growing season in the North, there appeared to be genetic differentiation in physiological integration within the Northern region. This shows that both regional and local conditions need to be taken into account in future studies of genetic differentiation of physiological integration in clonal plants.

Predator faunas past and present: quantifying the influence of waterborne cues in divergent ecotypes of the isopod Asellus aquaticus.

Waterborne chemical cues are an important source of information for many aquatic organisms, in particular when assessing the current risk of predation. The ability to use chemical cues to detect and respond to potential predators before an actual encounter can improve prey chances of survival. We investigated predator recognition and the impact of chemical cues on predator avoidance in the freshwater isopod Asellus aquaticus. This isopod has recently colonised a novel habitat and diverged into two distinct ecotypes, which encounter different predator communities. Using laboratory-based choice experiments, we have quantified behavioural responses to chemical cues from predators typical of the two predator communities (larval dragonflies in the ancestral habitat, perch in the newly colonised habitat) in wild-caught and lab-reared Asellus of the two ecotypes. Individuals with prior experience of predators showed strong predator avoidance to cues from both predator types. Both ecotypes showed similar antipredator responses, but sexes differed in terms of threat-sensitive responses with males avoiding areas containing predator cues to a larger extent than females. Overall, chemical cues from fish elicited stronger predator avoidance than cues from larval dragonflies. Our results indicate that in these isopods, prior exposure to predators is needed to develop antipredator behaviour based on waterborne cues. Furthermore, the results emphasise the need to analyse predator avoidance in relation to waterborne cues in a sex-specific context, because of potential differences between males and females in terms of vulnerability and life history strategies.

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.

Seasonal migration to high latitudes results in major reproductive benefits in an insect.

Little is known of the population dynamics of long-range insect migrants, and it has been suggested that the annual journeys of billions of nonhardy insects to exploit temperate zones during summer represent a sink from which future generations seldom return (the "Pied Piper" effect). We combine data from entomological radars and ground-based light traps to show that annual migrations are highly adaptive in the noctuid moth Autographa gamma (silver Y), a major agricultural pest. We estimate that 10-240 million immigrants reach the United Kingdom each spring, but that summer breeding results in a fourfold increase in the abundance of the subsequent generation of adults, all of which emigrate southward in the fall. Trajectory simulations show that 80% of emigrants will reach regions suitable for winter breeding in the Mediterranean Basin, for which our population dynamics model predicts a winter carrying capacity only 20% of that of northern Europe during the summer. We conclude not only that poleward insect migrations in spring result in major population increases, but also that the persistence of such species is dependent on summer breeding in high-latitude regions, which requires a fundamental change in our understanding of insect migration.

Long-term persistence of GM oilseed rape in the seedbank.

Coexistence between genetically modified (GM) and non-GM plants is a field of rapid development and considerable controversy. In crops, it is increasingly important to understand and predict the GM volunteer emergence in subsequent non-GM crops. Theoretical models suggest recruitment from the seedbank over extended periods, but empirical evidence matching these predictions has been scarce. Here, we provide evidence of long-term GM seed persistence in conventional agriculture. Ten years after a trial of GM herbicide-tolerant oilseed rape, emergent seedlings were collected and tested for herbicide tolerance. Seedlings that survived the glufosinate herbicide (15 out of 38 volunteers) tested positive for at least one GM insert. The resulting density was equivalent to 0.01 plants m-2, despite complying with volunteer reduction recommendations. These results are important in relation to debating and regulating coexistence of GM and non-GM crops, particularly for planting non-GM crops after GM crops in the same field.

Energetic consequences of an inducible morphological defence in crucian carp.

Crucian carp (Carassius carassius) increases in body depth in response to chemical cues from piscivores and the deeper body constitutes a morphological defence against gape-limited piscivores. In the field, deep-bodied individuals suffer a density-dependent cost when competing with shallow-bodied conspecifics. Here, we use hydrodynamic theory and swimming respirometry to investigate the proposed mechanism underlying this effect, high drag caused by the deep-bodied morphology. Our study confirms that drag is higher for deep-bodied crucian carp, both in terms of estimated theoretical drag and power curve steepness. However, deep-bodied fish swimming at the velocity associated with minimum cost of transport, U mc, did not experience higher costs of transport than shallow-bodied fish. Deep-bodied crucian carp had significantly lower standard metabolic rates, i.e. metabolic rates at rest, and also lower U mc, and the resulting costs of transport were similar for the two morphs. Nevertheless, when deep-bodied individuals deviate from U mc, e.g. when increasing foraging effort under competition, their steeper power curves will cause substantial energy costs relative to shallow-bodied conspecifics. Furthermore, there is evidence that reductions in standard metabolic rate incur costs in terms of lower stress tolerance, reduced growth rate, and life history changes. Thus, this work provides links between hydrodynamics, a cost-reducing mechanism, and a density-dependent fitness cost associated with an inducible defence.

Benefits of a predtor-induced morphology in crucian carp.

Crucian carp (Carassius carassius) develop a deeper body in response to chemical cues from piscivores. This change in body morphology has been suggested to be a predator-induced defence. Here we investigate the possible benefits of the induced body morphology in laboratory experiments. Pike foraging behaviour when feeding on crucian carp of different body depths was recorded using video. Further, in a preference experiment pike were allowed to choose between shallow-bodied and deep-bodied crucian carp of similar lengths. Crucian carp body morphology did not affect predatory behaviours (activity, searching, following, observing, capture success) in northern pike, but an increase in crucian carp body depth led to an increase in handling time in pike. In the preference experiment, pike preferred shallow-bodied crucian carp over deep-bodied. Thus, a change in body morphology, induced by the presence of piscivores, benefits crucian carp by increasing piscivore handling times and an avoidance of the deep-bodied phenotype.

Trading off safety against food: state dependent habitat choice and foraging in crucian carp.

The influence of hunger level and predation risk on habitat choice and foraging in crucian carp, Carassius carassius, were studied in a laboratory experiment. Experiments were carried out in aquaria with or without a predator (pike, Esox lucius). Habitat use and foraging activity of three-fish foraging groups of either fed or hungry crucian carp were studied. Fish were allowed to choose between an open (risky) habitat with Tubifex worms and a habitat with dense vegetation (safe) without food. Habitat use was significantly affected by both risk of predation and hunger level. Crucian carp spent less time in the open habitat when there was a predator present and they also spent less time there when fed than when hungry. Furthermore, there was a significant interaction between risk of predation and hunger level, indicating a state-dependent trade-off between food acquisition and predator avoidance.