Spring temperature drives phenotypic selection on plasticity of flowering time.

In seasonal environments, a high responsiveness of development to increasing temperatures in spring can infer benefits in terms of a longer growing season, but also costs in terms of an increased risk of facing unfavourable weather conditions. Still, we know little about how climatic conditions influence the optimal plastic response. Using 22 years of field observations for the perennial forest herb Lathyrus vernus, we assessed phenotypic selection on among-individual variation in reaction norms of flowering time to spring temperature, and examined if among-year variation in selection on plasticity was associated with spring temperature conditions. We found significant among-individual variation in mean flowering time and flowering time plasticity, and that plants that flowered earlier also had a more plastic flowering time. Selection favoured individuals with an earlier mean flowering time and a lower thermal plasticity of flowering time. Less plastic individuals were more strongly favoured in colder springs, indicating that spring temperature influenced optimal flowering time plasticity. Our results show how selection on plasticity can be linked to climatic conditions, and illustrate how we can understand and predict evolutionary responses of organisms to changing environmental conditions.

Life-history characteristics and historical factors are important to explain regional variation in reproductive traits and genetic diversity in perennial mosses.

BACKGROUND AND AIMS: Plants have evolved an unrivalled diversity of reproductive strategies, including variation in the degree of sexual vs. clonal reproduction. This variation has important effects on the dynamics and genetic structure of populations. We examined the association between large-scale variation in reproductive patterns and intraspecific genetic diversity in two moss species where sex is manifested in the dominant haploid generation and sex expression is irregular. We predicted that in regions with more frequent realized sexual reproduction, populations should display less skewed sex ratios, should more often express sex and should have higher genetic diversity than in regions with largely clonal reproduction. METHODS: We assessed reproductive status and phenotypic sex in the dioicous long-lived Drepanocladus trifarius and D. turgescens, in 248 and 438 samples across two regions in Scandinavia with frequent or rare realized sexual reproduction, respectively. In subsets of the samples, we analysed genetic diversity using nuclear and plastid sequence information and identified sex with a sex-specific molecular marker in non-reproductive samples. KEY RESULTS: Contrary to our predictions, sex ratios did not differ between regions; genetic diversity did not differ in D. trifarius and it was higher in the region with rare sexual reproduction in D. turgescens. Supporting our predictions, relatively more samples expressed sex in D. trifarius in the region with frequent sexual reproduction. Overall, samples were mostly female. The degree of sex expression and genetic diversity differed between sexes. CONCLUSIONS: Sex expression levels, regional sex ratios and genetic diversity were not directly associated with the regional frequency of realized sexual reproduction, and relationships and variation patterns differed between species. We conclude that a combination of species-specific life histories, such as longevity, overall degree of successful sexual reproduction and recruitment, and historical factors are important to explain this variation. Our data on haploid-dominated plants significantly complement plant reproductive biology.

Forest edge effects on moss growth are amplified by drought.

Forest fragmentation increases the amount of edges in the landscape. Differences in wind, radiation, and vegetation structure create edge-to-interior gradients in forest microclimate, and these gradients are likely to be more pronounced during droughts and heatwaves. Although the effects of climate extremes on edge influences have potentially strong and long-lasting impacts on forest understory biodiversity, they are not well understood and are not often considered in management and landscape planning. Here we used a novel method of retrospectively quantifying growth to assess biologically relevant edge influences likely caused by microclimate using Hylocomium splendens, a moss with annual segments. We examined how spatio-temporal variation in drought across 3 years and 46 sites in central Sweden, affected the depth and magnitude of edge influences. We also investigated whether edge effects during drought were influenced by differences in forest structure. Edge effects were almost twice as strong in the drought year compared to the non-drought years, but we did not find clear evidence that they penetrated deeper into the forest in the drought year. Edge influences were also greater in areas that had fewer days with rain during the drought year. Higher levels of forest canopy cover and tree height buffered the magnitude of edge influence in times of drought. Our results demonstrate that edge effects are amplified by drought, suggesting that fragmentation effects are aggravated when droughts become more frequent and severe. Our results suggest that dense edges and buffer zones with high canopy cover can be important ways to mitigate negative drought impacts in forest edges.

Biotic and anthropogenic forces rival climatic/abiotic factors in determining global plant population growth and fitness.

Multiple, simultaneous environmental changes, in climatic/abiotic factors, interacting species, and direct human influences, are impacting natural populations and thus biodiversity, ecosystem services, and evolutionary trajectories. Determining whether the magnitudes of the population impacts of abiotic, biotic, and anthropogenic drivers differ, accounting for their direct effects and effects mediated through other drivers, would allow us to better predict population fates and design mitigation strategies. We compiled 644 paired values of the population growth rate (λ) from high and low levels of an identified driver from demographic studies of terrestrial plants. Among abiotic drivers, natural disturbance (not climate), and among biotic drivers, interactions with neighboring plants had the strongest effects on λ However, when drivers were combined into the 3 main types, their average effects on λ did not differ. For the subset of studies that measured both the average and variability of the driver, λ was marginally more sensitive to 1 SD of change in abiotic drivers relative to biotic drivers, but sensitivity to biotic drivers was still substantial. Similar impact magnitudes for abiotic/biotic/anthropogenic drivers hold for plants of different growth forms, for different latitudinal zones, and for biomes characterized by harsher or milder abiotic conditions, suggesting that all 3 drivers have equivalent impacts across a variety of contexts. Thus, the best available information about the integrated effects of drivers on all demographic rates provides no justification for ignoring drivers of any of these 3 types when projecting ecological and evolutionary responses of populations and of biodiversity to environmental changes.

Global gene flow releases invasive plants from environmental constraints on genetic diversity.

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area.

The relationship between pathogen life-history traits and metapopulation dynamics.

Plant pathogen traits, such as transmission mode and overwintering strategy, may have important effects on dispersal and persistence, and drive disease dynamics. Still, we lack insights into how life-history traits influence spatiotemporal disease dynamics. We adopted a multifaceted approach, combining experimental assays, theory and field surveys, to investigate whether information about two pathogen life-history traits - infectivity and overwintering strategy - can predict pathogen metapopulation dynamics in natural systems. For this, we focused on four fungal pathogens (two rust fungi, one chytrid fungus and one smut fungus) on the forest herb Anemone nemorosa. Pathogens infecting new plants mostly via spores (the chytrid and smut fungi) had higher patch occupancies and colonization rates than pathogens causing mainly systemic infections and overwintering in the rhizomes (the two rust fungi). Although the rust fungi more often occupied well-connected plant patches, the chytrid and smut fungi were equally or more common in isolated patches. Host patch size was positively related to patch occupancy and colonization rates for all pathogens. Predicting disease dynamics is crucial for understanding the ecological and evolutionary dynamics of host-pathogen interactions, and to prevent disease outbreaks. Our study shows that combining experiments, theory and field observations is a useful way to predict disease dynamics.

Spring and autumn phenology in an understory herb are uncorrelated and driven by different factors.

PREMISE: Climate warming has altered the start and end of growing seasons in temperate regions. Ultimately, these changes occur at the individual level, but little is known about how previous seasonal life-history events, temperature, and plant-resource state simultaneously influence the spring and autumn phenology of plant individuals. METHODS: We studied the relationships between the timing of leaf-out and shoot senescence over 3 years in a natural population of the long-lived understory herb Lathyrus vernus and investigated the effects of spring temperature, plant size, reproductive status, and grazing on spring and autumn phenology. RESULTS: The timing of leaf-out and senescence were consistent within individuals among years. Leaf-out and senescence were not correlated with each other within years. Larger plants leafed out and senesced later, and size had no effect on growing season length. Reproductive plants leafed out earlier and had longer growing seasons than nonreproductive plants. Grazing had no detectable effects on phenology. Colder spring temperatures delayed senescence in two of three study years. CONCLUSIONS: The timing of seasonal events, such as leaf-out and senescence in plants can be expressed largely independently within and among seasons and are influenced by different factors. Growing season start and length can often be dependent on plant condition and reproductive status. Knowledge about the drivers of growing season length of individuals is essential to more accurately predict species and community responses to environmental variation.

Selection against early flowering in geothermally heated soils is associated with pollen but not prey availability in a carnivorous plant.

PREMISE: In high-latitude environments, plastic responses of phenology to increasing spring temperatures allow plants to extend growing seasons while avoiding late frosts. However, evolved plasticity might become maladaptive if climatic conditions change and spring temperatures no longer provide reliable cues for conditions important for fitness. Maladaptative phenological responses might be related to both abiotic factors and mismatches with interacting species. When mismatches arise, we expect selection to favor changes in phenology. METHODS: We combined observations along a soil temperature gradient in a geothermally heated area with pollen and prey supplementation experiments and examined how phenotypic selection on flowering time in the carnivorous plant Pinguicula vulgaris depends on soil temperature, and pollen and prey availability. RESULTS: Flowering advanced and fitness decreased with increasing soil temperature. However, in pollen-supplemented plants, fitness instead increased with soil temperature. In heated soils, there was selection favoring later flowering, while earlier flowering was favored in unheated soils. This pattern remained also after artificially increasing pollen and prey availability. CONCLUSIONS: Plant-pollinator mismatches can be an important reason why evolved plastic responses of flowering time to increasing spring temperatures become maladaptive under novel environmental conditions, and why there is selection to delay flowering. In our study, selection for later flowering remained after artificially increasing pollen availability, suggesting that abiotic factors also contribute to the observed selection. Identifying the factors that make evolved phenological responses maladaptive under novel conditions is fundamental for understanding and predicting evolutionary responses to climate warming.

Phenotypic plasticity masks range-wide genetic differentiation for vegetative but not reproductive traits in a short-lived plant.

Genetic differentiation and phenotypic plasticity jointly shape intraspecific trait variation, but their roles differ among traits. In short-lived plants, reproductive traits may be more genetically determined due to their impact on fitness, whereas vegetative traits may show higher plasticity to buffer short-term perturbations. Combining a multi-treatment greenhouse experiment with observational field data throughout the range of a widespread short-lived herb, Plantago lanceolata, we (1) disentangled genetic and plastic responses of functional traits to a set of environmental drivers and (2) assessed how genetic differentiation and plasticity shape observational trait-environment relationships. Reproductive traits showed distinct genetic differentiation that largely determined observational patterns, but only when correcting traits for differences in biomass. Vegetative traits showed higher plasticity and opposite genetic and plastic responses, masking the genetic component underlying field-observed trait variation. Our study suggests that genetic differentiation may be inferred from observational data only for the traits most closely related to fitness.

Pathogen infection influences the relationship between spring and autumn phenology at the seedling and leaf level.

Seasonal life history events are often interdependent, but we know relatively little about how the relationship between different events is influenced by the abiotic and biotic environment. Such knowledge is important for predicting the immediate and evolutionary phenological response of populations to changing conditions. We manipulated germination timing and shade in a multi-factorial experiment to investigate the relationship between spring and autumn phenology in seedlings of the pedunculate oak, Quercus robur, and whether this relationship was mediated by natural colonization of leaves by specialist fungal pathogens (i.e., the oak powdery mildew complex). Each week delay in germination corresponded to about 2 days delay in autumn leaf senescence, and heavily shaded seedlings senesced 5-8 days later than seedlings in light shade or full sun. Within seedlings, leaves on primary-growth shoots senesced later than those on secondary-growth shoots in some treatments. Path analyses demonstrated that germination timing and shade affected autumn phenology both directly and indirectly via pathogen load, though the specific pattern differed among and within seedlings. Pathogen load increased with later germination and greater shade. Greater pathogen load was in turn associated with later senescence for seedlings, but with earlier senescence for individual leaves. Our findings show that relationships between seasonal events can be partly mediated by the biotic environment and suggest that these relationships may differ between the plant and leaf level. The influence of biotic interactions on phenological correlations across scales has implications for understanding phenotypic variation in phenology and for predicting how populations will respond to climatic perturbation.

Global shifts in the phenological synchrony of species interactions over recent decades.

Phenological responses to climate change (e.g., earlier leaf-out or egg hatch date) are now well documented and clearly linked to rising temperatures in recent decades. Such shifts in the phenologies of interacting species may lead to shifts in their synchrony, with cascading community and ecosystem consequences. To date, single-system studies have provided no clear picture, either finding synchrony shifts may be extremely prevalent [Mayor SJ, et al. (2017) Sci Rep 7:1902] or relatively uncommon [Iler AM, et al. (2013) Glob Chang Biol 19:2348-2359], suggesting that shifts toward asynchrony may be infrequent. A meta-analytic approach would provide insights into global trends and how they are linked to climate change. We compared phenological shifts among pairwise species interactions (e.g., predator-prey) using published long-term time-series data of phenological events from aquatic and terrestrial ecosystems across four continents since 1951 to determine whether recent climate change has led to overall shifts in synchrony. We show that the relative timing of key life cycle events of interacting species has changed significantly over the past 35 years. Further, by comparing the period before major climate change (pre-1980s) and after, we show that estimated changes in phenology and synchrony are greater in recent decades. However, there has been no consistent trend in the direction of these changes. Our findings show that there have been shifts in the timing of interacting species in recent decades; the next challenges are to improve our ability to predict the direction of change and understand the full consequences for communities and ecosystems.

Climate drives among-year variation in natural selection on flowering time.

To predict long-term responses to climate change, we need to understand how changes in temperature and precipitation elicit both immediate phenotypic responses and changes in natural selection. We used 22 years of data for the perennial herb Lathyrus vernus to examine how climate influences flowering phenology and phenotypic selection on phenology. Plants flowered earlier in springs with higher temperatures and higher precipitation. Early flowering was associated with a higher fitness in nearly all years, but selection for early flowering was significantly stronger in springs with higher temperatures and lower precipitation. Climate influenced selection through trait distributions, mean fitness and trait-fitness relationships, the latter accounting for most of the among-year variation in selection. Our results show that climate both induces phenotypic responses and alters natural selection, and that the change in the optimal phenotype might be either weaker, as for spring temperature, or stronger, as for precipitation, than the optimal response.

The timing and asymmetry of plant-pathogen-insect interactions.

Insects and pathogens frequently exploit the same host plant and can potentially impact each other's performance. However, studies on plant-pathogen-insect interactions have mainly focused on a fixed temporal setting or on a single interaction partner. In this study, we assessed the impact of time of attacker arrival on the outcome and symmetry of interactions between aphids (Tuberculatus annulatus), powdery mildew (Erysiphe alphitoides), and caterpillars (Phalera bucephala) feeding on pedunculate oak, Quercus robur, and explored how single versus multiple attackers affect oak performance. We used a multifactorial greenhouse experiment in which oak seedlings were infected with either zero, one, two, or three attackers, with the order of attacker arrival differing among treatments. The performances of all involved organisms were monitored throughout the experiment. Overall, attackers had a weak and inconsistent impact on plant performance. Interactions between attackers, when present, were asymmetric. For example, aphids performed worse, but powdery mildew performed better, when co-occurring. Order of arrival strongly affected the outcome of interactions, and early attackers modified the strength and direction of interactions between later-arriving attackers. Our study shows that interactions between plant attackers can be asymmetric, time-dependent, and species specific. This is likely to shape the ecology and evolution of plant-pathogen-insect interactions.

Hiding from the climate: Characterizing microrefugia for boreal forest understory species.

Climate warming is likely to shift the range margins of species poleward, but fine-scale temperature differences near the ground (microclimates) may modify these range shifts. For example, cold-adapted species may survive in microrefugia when the climate gets warmer. However, it is still largely unknown to what extent cold microclimates govern the local persistence of populations at their warm range margin. We located 99 microrefugia, defined as sites with edge populations of 12 widespread boreal forest understory species (vascular plants, mosses, liverworts and lichens) in an area of ca. 24,000 km2 along the species' southern range margin in central Sweden. Within each population, a logger measured temperature eight times per day during one full year. Using univariate and multivariate analyses, we examined the differences of the populations' microclimates with the mean and range of microclimates in the landscape, and identified the typical climate, vegetation and topographic features of these habitats. Comparison sites were drawn from another logger data set (n = 110), and from high-resolution microclimate maps. The microrefugia were mainly places characterized by lower summer and autumn maximum temperatures, late snow melt dates and high climate stability. Microrefugia also had higher forest basal area and lower solar radiation in spring and autumn than the landscape average. Although there were common trends across northern species in how microrefugia differed from the landscape average, there were also interspecific differences and some species contributed more than others to the overall results. Our findings provide biologically meaningful criteria to locate and spatially predict potential climate microrefugia in the boreal forest. This opens up the opportunity to protect valuable sites, and adapt forest management, for example, by keeping old-growth forests at topographically shaded sites. These measures may help to mitigate the loss of genetic and species diversity caused by rear-edge contractions in a warmer climate.

Diversity of ageing across the tree of life.

Evolution drives, and is driven by, demography. A genotype moulds its phenotype's age patterns of mortality and fertility in an environment; these two patterns in turn determine the genotype's fitness in that environment. Hence, to understand the evolution of ageing, age patterns of mortality and reproduction need to be compared for species across the tree of life. However, few studies have done so and only for a limited range of taxa. Here we contrast standardized patterns over age for 11 mammals, 12 other vertebrates, 10 invertebrates, 12 vascular plants and a green alga. Although it has been predicted that evolution should inevitably lead to increasing mortality and declining fertility with age after maturity, there is great variation among these species, including increasing, constant, decreasing, humped and bowed trajectories for both long- and short-lived species. This diversity challenges theoreticians to develop broader perspectives on the evolution of ageing and empiricists to study the demography of more species.

A natural heating experiment: Phenotypic and genotypic responses of plant phenology to geothermal soil warming.

Under global warming, the survival of many populations of sedentary organisms in seasonal environments will largely depend on their ability to cope with warming in situ by means of phenotypic plasticity or adaptive evolution. This is particularly true in high-latitude environments, where current growing seasons are short, and expected temperature increases large. In such short-growing season environments, the timing of growth and reproduction is critical to survival. Here, we use the unique setting provided by a natural geothermal soil warming gradient (Hengill geothermal area, Iceland) to study the response of Cerastium fontanum flowering phenology to temperature. We hypothesized that trait expression and phenotypic selection on flowering phenology are related to soil temperature, and tested the hypothesis that temperature-driven differences in selection on phenology have resulted in genetic differentiation using a common garden experiment. In the field, phenology was related to soil temperature, with plants in warmer microsites flowering earlier than plants at colder microsites. In the common garden, plants responded to spring warming in a counter-gradient fashion; plants originating from warmer microsites flowered relatively later than those originating from colder microsites. A likely explanation for this pattern is that plants from colder microsites have been selected to compensate for the shorter growing season by starting development at lower temperatures. However, in our study we did not find evidence of variation in phenotypic selection on phenology in relation to temperature, but selection consistently favoured early flowering. Our results show that soil temperature influences trait expression and suggest the existence of genetically based variation in flowering phenology leading to counter-gradient local adaptation along a gradient of soil temperatures. An important implication of our results is that observed phenotypic responses of phenology to global warming might often be a combination of short-term plastic responses and long-term evolutionary responses, acting in different directions.

Resource overlap and dilution effects shape host plant use in a myrmecophilous butterfly.

The effects of consumers on fitness of resource organisms are a complex function of the spatio-temporal distribution of the resources, consumer functional responses and trait preferences, and availability of other resources. The ubiquitous variation in the intensity of species interactions has important consequences for the ecological and evolutionary dynamics of natural populations. Nevertheless, little is known about the processes causing this variation and their operational scales. Here, we examine how variation in the intensity of a consumer-resource interaction is related to resource timing, resource density and abundance of other resources. Using the butterfly consumer Phengaris alcon and its two sequential resources, the host plant Gentiana pneumonanthe and the host ants Myrmica spp., we investigated how butterfly egg-laying depended on focal host plant phenology, density and phenology of neighbouring host plants and host ant abundance. Butterflies preferred plants that simultaneously maximized the availability of both larval resources in time and space, that is, they chose early-flowering plants that were of higher nutritional quality for larvae where host ants were abundant. Both the probability of oviposition and the number of eggs were lower in plant individuals with a high neighbour density than in more isolated plants, and this dilution effect was stronger when neighbours flowered early. Our results show that plant-herbivore interactions simultaneously depend on the spatio-temporal distribution of a focal resource and on the small-scale spatial variation in the abundance of other herbivore resources. Given that consumers have negative effects on fitness and prefer certain timing of the resource organisms, this implies that processes acting at the levels of individuals, populations and communities simultaneously contribute to variation in consumer-mediated natural selection.

Sex and the cost of reproduction through the life course of an extremely long-lived herb.

Despite being central concepts for life history theory, little is known about how reproductive effort and costs vary with individual age once plants have started to reproduce. We conducted a 5-year field study and estimated age-dependent reproductive effort for both sexes in the extraordinarily long-lived dioecious plant Borderea pyrenaica. We also evaluated costs of reproduction on vital rates for male and female plants, both by examining effects of differences in individual reproductive effort under natural conditions, and by conducting a flower removal experiment, aimed at decreasing reproductive effort. Reproductive effort was fairly constant and independent of age for males, which may reflect a strategy of adjusting overall reproductive output by spreading reproduction over the life course. Females had a higher total effort, which first increased and then decreased with age. The latter may be a response to an increasing reproductive value-an inverse of a terminal investment-or a sign of reproductive senescence due to an age-related physiological decline. Seed production was lower in plants with higher previous reproductive effort and this effect increased with age. We found no evidence for costs of reproduction on other vital rates for either sex. Experimental flower removal only resulted in progressively more negative effects on flower production in older male plants, whereas female vital rates were unaffected. Overall, this study demonstrates that not only sex, but also age influences resource allocation trade-offs and, thus, plant life history evolution.

Grazers affect selection on inflorescence height both directly and indirectly and effects change over time.

Selection mediated by one biotic agent will often be modified by the presence of other biotic interactions, and the importance of such indirect effects might change over time. We conducted an 11-yr field experiment to test the prediction that large grazers affect selection on floral display of the dimorphic herb Primula farinosa not only directly through differential grazing damage, but also indirectly by affecting vegetation height and thereby selection mediated by pollinators and seed predators. Exclusion of large grazers increased vegetation height and the strength of pollinator-mediated selection for tall inflorescences and seed-predator-mediated selection for short inflorescences. The direct effect of grazers on selection resulting from differential grazing damage to the two scape morphs showed no temporal trend. By contrast, the increase in vegetation height in exclosures over time was associated with an increase in selection mediated by pollinators and seed predators. In the early years of the experiment, the indirect effects of grazers on selection mediated by pollinators and seed predators were weak, whereas at the end of the experiment, the indirect effects were of similar magnitude as the direct effect due to differential grazing damage. The results demonstrate that the indirect effects of a selective agent can be as strong as its direct effects, and that the relative importance of direct vs. indirect effects on selection can change over time. A full understanding of the ecological processes governing variation in selection thus requires that both direct and indirect effects of biotic interactions are assessed.

Phenological synchrony between a butterfly and its host plants: Experimental test of effects of spring temperature.

Climate-driven changes in the relative phenologies of interacting species may potentially alter the outcome of species interactions. Phenotypic plasticity is expected to be important for short-term response to new climate conditions, and differences between species in plasticity are likely to influence their temporal overlap and interaction patterns. As reaction norms of interacting species may be locally adapted, any such climate-induced change in interaction patterns may vary among localities. However, consequences of spatial variation in plastic responses for species interactions are understudied. We experimentally explored how temperature affected synchrony between spring emergence of a butterfly, Anthocharis cardamines, and onset of flowering of five of its host plant species across a latitudinal gradient. We also studied potential effects on synchrony if climate-driven northward expansions would be faster in the butterflies than in host plants. Lastly, to assess how changes in synchrony influence host use we carried out an experiment to examine the importance of the developmental stage of plant reproductive structures for butterfly oviposition preference. In southern locations, the butterflies were well-synchronized with the majority of their local host plant species across temperatures, suggesting that thermal plasticity in butterfly development matches oviposition to host plant development and that thermal reaction norms of insects and plants result in similar advancement of spring phenology in response to warming. In the most northern region, however, relative phenology between the butterfly and two of its host plant species changed with increased temperature. We also show that the developmental stage of plants was important for egg-laying, and conclude that temperature-induced changes in synchrony in the northernmost region are likely to lead to shifts in host use in A. cardamines if spring temperatures become warmer. Northern expansion of butterfly populations might possibly have a positive effect on keeping up with host plant phenology with more northern host plant populations. Considering that the majority of insect herbivores exploit multiple plant species differing in their phenological response to spring temperatures, temperature-induced changes in synchrony might lead to shifts in host use and changes in species interactions in many temperate communities.