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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.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.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.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.

[Aim]: Range shifts are expected to occur when populations at one range margin per-form better than those at the other margin, yet no global trend in population per-formances at range margins has been demonstrated empirically across a wide range of taxa and biomes. Here we test the prediction that, if impacts of ongoing climate change on performance in marginal populations are widespread, then populations from the high- latitude margin (HLM) should perform as well as or better than central populations, whereas low- latitude margin (LLM) populations should perform worse., [Location]: Global., [Time period]: 1995–2019., [Major taxa studied]: Plants and animals., [Methods]: To test our prediction, we used a meta-analysis to quantify empirical sup-port for asymmetry in the performance of high- and low- latitude margin populations compared to central populations. Performance estimates (survival, reproduction, or lifetime fitness) for populations occurring in their natural environment were derived from 51 papers involving 113 margin-centre comparisons from 54 species and 705 populations from the Americas, Europe, Africa and Australia. We then related these performance differences to climatic differences among populations. We also tested whether patterns are consistent across taxonomic kingdoms (plants vs animals) and across realms (marine vs terrestrial)., [Results]: Populations at margins performed significantly worse than central popula-tions, and this trend was primarily driven by the low- latitude margin. Although the dif-ference was of small magnitude, it was largely consistent across biological kingdoms and realms. Differences in performance were weakly (p= .08) related to the differ-ence in average temperatures between central and marginal populations., [Main conclusions]: The observed asymmetry in performance in marginal populations is consistent with predictions about the effects of global climate change, though fur-ther research is needed to confirm the effect of climate. It indicates that changes in demographic rates in marginal populations can serve as early-warning signals of impending range shifts, This study was funded by Nord Forsk grant no. 80167 to the NORA Consortium (Nordic Network for the Study of Species Range Dynamics, 2009–2012), by projects POPULIM (CGL2010-22180) and PERSLIM (CGL2010-18381) of the Spanish Ministerio de Ciencia e Innovación, the EU ERA- net BiodivERsA project BeFoFu (via funding to ASJ by Natural Environmental Research Council grant NE/G002118/1) and the Institut National de Recherche pour l’agriculture, l’alimentation et l’environnement Assessing Climate Change Adaptation Framework project FORADAPT. FRS was funded by a postdoctoral fellow-ship from the Spanish Ministerio de Economía y Competitividad (FPD2013-16756) and a Talent Attraction grant from the VI Plan Propio de Investigación at Universidad de Sevilla (VIPPIT-2018- IV.2). JCS considers this work a contribution to his VILLUM Investigator project ‘Biodiversity Dynamics in a Changing World’, funded by VILLUM FONDEN (grant 16549), and Center for Ecological Dynamics in a Novel Biosphere (ECONOVO), funded by Danish National Research Foundation (grant DNRF173).

The local climate in forest understories can deviate substantially from ambient conditions. Moreover, forest microclimates are often characterized by cyclic changes driven by management activities such as clear-cutting and subsequent planting. To understand how and why understorey plant communities change, both ambient climate change and temporal variation in forest structure have to be considered. We used inventories from 11,436 productive forest sites in Sweden repeated every 10th year 1993–2017 to examine how variation in forest structure influences changes in the average value of minimum and maximum temperature preferences of all species in a community, that is, community temperature indices (CTIs). We then evaluated to what extent these changes were driven by local extinctions and colonizations, respectively, and to what extent the difference in CTI value between two inventories was related to changes in forest density and in macroclimate. Lastly, we tested whether effects on CTI change by these two drivers were modified by topography, soil moisture and tree species composition. CTI values of the understorey plant communities increased after clear-cutting, and decreased during periods when the forest grew denser. During the period immediately after clear-cutting, changes were predominately driven by colonizations of species with a preference for higher temperatures. During the forest regeneration phase, both colonizations by species preferring lower temperatures and local extinctions of species preferring higher temperatures increased. The change in understorey CTI over 10-year periods was explained more by changes in forest density, than by changes in macroclimate. Soil moisture, topography and forest tree species composition modified to some extent the effects of changes in forest density and in macroclimate on understorey CTI values. Synthesis. Via stand manipulation, forest management impacts the effects of regional climate on understorey plant communities. This implies that forest management by creating denser stands locally even can counterbalance the effects of regional changes in climate. Consequently, interpretations of changes in the mean temperature preference of species in forest understorey communities should take forest management regimes into account.

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.

Species and community-level responses to warming are well documented, with plants and invertebrates known to alter their range, phenology or composition as temperature increases. The effects of warming on biotic interactions are less clearly understood, but can have consequences that cascade through ecological networks. Here, we used a natural soil temperature gradient of 5–35°C in the Hengill geothermal valley, Iceland, to investigate the effects of temperature on plant community composition and plant–invertebrate interactions. We quantified the level of invertebrate herbivory on the plant community across the temperature gradient and the interactive effects of temperature, plant phenology (i.e. development stage) and vegetation community composition on the probability of herbivory for three ubiquitous plant species, Cardamine pratensis, Cerastium fontanum and Viola palustris. We found that the percentage cover of graminoids and forbs increased, while the amount of litter decreased, with increasing soil temperature. Invertebrate herbivory also increased with soil temperature at the plant community level, but this was underpinned by different effects of temperature on herbivory for individual plant species, mediated by the seasonal development of plants and the composition of the surrounding vegetation. This illustrates the importance of considering the development stage of organisms in climate change research given the variable effects of temperature on susceptibility to herbivory at different ontogenetic stages.

1. The histories of individuals impact the dynamics of their populations. Matrix projection models (MPMs) are used to analyse population dynamics, but are not structured to incorporate these influe ...

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.

Increased anthropogenic influence on the environment has accentuated the need to assess how climate and other environmental factors drive vital rates and population dynamics of different types of organisms. However, to allow distinction between the effects of multiple correlated variables, and to capture the effects of rare and extreme climatic conditions, studies extending over decades are often necessary. In this study, we used an individual-based dataset collected in three populations of Pulsatilla vulgaris subsp. gotlandica during 34 years, to explore the effects of variation in precipitation and temperature on vital rates and population dynamics. Most of the observed conspicuous variation in flowering among years was associated with differences in precipitation and temperature in the previous summer and autumn with a higher incidence of flowering following summers with high precipitation and low temperatures. In contrast, climatic variables had no significant effects on individual growth or survival. Although the weather-driven variation in flowering had only moderate absolute effects on the population growth rate, simulated persistent changes in average precipitation and temperature resulted in considerable reductions in population sizes compared with current conditions. Analyses carried out with subsets of data consisting of 5 and 10 years yielded results that strongly deviated from those based on the full dataset. Synthesis. The results of this study illustrate the importance of long-term demographic monitoring to identify key climatic variables affecting vital rates and driving population dynamics in long-lived organisms.

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 three 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.KEY 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 both leafed out and senesced later, and there was no effect of size on growing season length. Reproductive plants leafed out earlier and had longer growing seasons than non-reproductive plants. Grazing had no detectable effects on phenology. Colder spring temperatures delayed senescence in two of three study years.CONCLUSION: 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 condition-dependent and dependent on plant reproductive status. To more accurately predict species and community responses to environmental variation, knowledge about the drivers of growing season length of individuals is essential. This article is protected by copyright. All rights reserved.

The timing of different life history events are often correlated, and selection might only rarely be exerted independently on the timing of a single event. In plants, phenotypic selection has often been shown to favour earlier flowering. However, little is known about to what extent this selection acts directly vs. indirectly via vegetative phenology, and if selection on the two traits is correlational. We estimated direct, indirect and correlational phenotypic selection on vegetative and reproductive phenology over three years for the perennial herb Lathyrus vernus. Direct selection favoured earlier flowering and shorter timespans between leaf-out and flowering in all years. However, early flowering was associated with early leaf-out, and the direction of selection on leaf-out day varied among years. As a result, selection on leaf-out weakened selection for early flowering in one of the study years. We found no evidence of correlational selection. Our results highlight the importance of including temporally correlated traits when exploring selection on the phenology of seasonal events.

Species at their warm range margin are potentially threatened by higher temperatures, but may persist in microrefugia. Whether such microsites occur due to more suitable microclimate or due to lower biotic pressure from, for example competitive species, is still not fully resolved. We examined whether boreal bryophytes and lichens show signs of direct climate limitation, that is whether they perform better in cold and/or humid microclimates at their warm range margin. We transplanted a moss, a liverwort and a lichen to 58 boreal forest sites with different microclimates at the species' southern range margin in central Sweden. Species were grown in garden soil patches to control the effects of competitive exclusion and soil quality. We followed the transplanted species over three growing seasons (2016–2018) and modelled growth and vitality for each species as a function of subcanopy temperature, soil moisture, air humidity and forest type. In 2018, we also recorded the cover of other plants having recolonized the garden soil patches and modelled this potential future competition with the same environmental variables plus litter. Species performance increased with warmer temperatures, which was often conditional on high soil moisture, and at sites with more conifers. Soil moisture had a positive effect, especially on the moss in the last year 2018, when the growing season was exceptionally hot and dry. The lichen was mostly affected by gastropod grazing. Recolonization of other plants was also faster at warmer and moister sites. The results indicate that competition, herbivory, shading leaf litter and water scarcity might be more important than the direct effects of temperature for performance at the species' warm range margin. Synthesis. In a transplant experiment with three boreal understorey species, we did not find signs of direct temperature limitation towards the south. Forest microrefugia, that is habitats where these species could persist regional warming, may instead be sites with fewer competitors and enemies, and with sufficient moisture and more conifers in the overstorey. Royal Swedish Academy of Sciences 2014-00530

Sex ratio variation is common among organisms with separate sexes. In bryophytes, sex chromosome segregation at meiosis suggests a balanced progeny sex ratio. However, most bryophyte populations exhibit female-biased phenotypic sex ratios based on the presence of reproductive structures on gametophytes. Many bryophyte populations do not form sexual organs, and genotypic sex ratio variation in such populations is mostly unknown. We tested sex expression, and phenotypic and genotypic sex ratios against environmental parameters in natural populations of the unisexual wetland moss Drepanocladus lycopodiodes at 11 sites in each of three regions in southern Sweden. We identified sex in 660 individual ramets, based on sexual structures, when present, or with a specifically designed molecular marker, when absent. All regions exhibited a female bias in phenotypic and genotypic sex ratios. Sex ratio biases and sex expression differed between regions. Sex ratios were less female-biased in larger patches. Wetter patches exhibited a stronger female bias in genotypic sex ratio and lower sex expression. This is the first evidence of environmental effects on genotypic sex ratio in mosses. A higher frequency of females in wet patches could be due to higher female resource demands for sporophyte production or higher male sensitivity to wetness. A higher incidence of females than males in moister sites aligns with female flowering plants, but differs from reproductive bryophytes in drier environments. Taken together with previous results, our data indicate that sex ratio variation and its drivers differ among species, their life histories and environments.

Selection on flowering time in plants is often mediated by multiple agents, including climatic conditions and the intensity of mutualistic and antagonistic interactions with animals. These selective agents can have both direct and indirect effects. For example, climate might not only influence phenotypic selection on flowering time directly by affecting plant physiology, but it can also alter selection indirectly by modifying the seasonal activity and relative timing of animals interacting with plants. We used 21 years of data to identify the drivers of selection on flowering time in the perennial herb Lathyrus vernus, and to examine if antagonistic plant-animal interactions mediate effects of climate on selection. We examined the fitness consequences of vertebrate grazing and pre-dispersal seed predation, and how these effects varied among years and among individuals within years. Although both antagonistic plant-animal interactions had important negative effects on plant fitness, only grazing intensity was consistently related to plant phenology, being higher in early-flowering individuals. Spring temperature influenced the intensity of both plant-animal interactions, as well as the covariance between seed predation and plant phenology. However, only differences in grazing intensity among years were associated with differences in selection on flowering time; the strength of selection for early flowering being stronger in years with lower mean intensity of grazing. Our results illustrate how climatic conditions can influence plant-animal interactions that are important selective agents for plant traits. A broader implication of our findings is that both ecological and evolutionary responses to climatic changes might be indirect, and largely mediated by species interactions.

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.

To understand how the environment drives spatial variation in population dynamics, we need to assess the effects of a large number of potential drivers on the vital rates (survival, growth and reproduction), and explore these relationships over large geographical areas and long environmental gradients. In this study, we examined the effects of a broad variety of abiotic and biotic environmental factors, including intraspecific density, on the demography of the forest understory herbActaea spicatabetween 2017 and 2019 at 40 sites across Sweden, including the northern range margin of its distribution. We assessed the effect of potential environmental drivers on vital rates using generalized linear mixed models (GLMMs), and then quantified the impact of each important driver on population growth rate (λ) using integral projection models (IPMs). Population dynamics ofA. spicatawere mostly driven by environmental factors affecting survival and growth, such as air humidity, soil depth and forest tree species composition, and thus those drivers jointly determined the realized niche of the species. Soil pH had a strong effect on the flowering probability, while the effect on population growth rate was relatively small. In addition to identifying specific drivers forA. spicata’spopulation dynamics, our study illustrates the impact that spatial variation in environmental conditions can have on λ. Assessing the effects of a broad range of potential drivers, as done in this study, is important not only to quantify the relative importance of different drivers for population dynamics but also to understand species distributions and abundance patterns.

Variation in the degree of synchrony among host plants and herbivores can disrupt or intensify species interactions, alter the strength of natural selection on traits associated with phenological timing, and drive novel host plant associations. We used field observations from three regions during four seasons to examine how timing of the butterfly herbivore Anthocharis cardamines relative to six host plant species (Arabis hirsuta, Cardamine pratensis, Arabis glabra, Arabidopsis thaliana, Thlaspi caerulescens and Capsella bursa-pastoris) influenced host species use and the choice of host plant individuals within populations. Butterflies laid a larger fraction of their eggs on species that were closer to the butterfly's preferred stage of development than on other host species. Within host plant populations, butterflies showed a stronger preference for individuals with a late phenology when plants within the population were on average more developed at the time of butterfly flight. Our results suggest that changes in synchrony between herbivores and their host plants are associated with changes in both host species use and the choice of host plant individuals differing in phenology within populations. This is likely to be an important mechanism generating variation in interaction intensities and trait selection in the wild, and therefore also relevant for understanding how anthropogenic induced changes, such as global warming, will influence natural communities., This study was funded by the Swedish Research Council (JE) and Séneca Foundation (postdoctoral grant 12337/PD/09 to JANC) for funding.

Variation in the intensity of plant-animal interactions over different spatial scales is widespread and might strongly influence fitness and trait selection in plants. Differences in traits among p ...

Temporal variation in natural selection has profound effects on the evolutionary trajectories of populations. One potential source of variation in selection is that differences in thermal reaction norms and temperature influence the relative phenology of interacting species. We manipulated the phenology of the butterfly herbivore Anthocharis cardamines relative to genetically identical populations of its host plant, Cardamine pratensis, and examined the effects on butterfly preferences and selection acting on the host plant. We found that butterflies preferred plants at an intermediate flowering stage, regardless of the timing of butterfly flight relative to flowering onset of the population. Consequently, the probability that plant genotypes differing in timing of flowering should experience a butterfly attack depended strongly on relative phenology. These results suggest that differences in spring temperature influence the direction of herbivore-mediated selection on flowering phenology, and that climatic conditions can influence natural selection also when phenotypic preferences remain constant.

International audience; The role of climate in determining range margins is often studied using species distribution models (SDMs), which are easily applied but have well-known limitations, e.g. due to their correlative nature and colonization and extinction time lags. Transplant experiments can give more direct information on environmental effects, but often cover small spatial and temporal scales. We simultaneously applied a SDM using high-resolution spatial predictors and an integral projection (demographic) model based on a transplant experiment at 58 sites to examine the effects of microclimate, light and soil conditions on the distribution and performance of a forest herb, Lathyrus vernus, at its cold range margin in central Sweden. In the SDM, occurrences were strongly associated with warmer climates. In contrast, only weak effects of climate were detected in the transplant experiment, whereas effects of soil conditions and light dominated. The higher contribution of climate in the SDM is likely a result from its correlation with soil quality, forest type and potentially historic land use, which were unaccounted for in the model. Predicted habitat suitability and population growth rate, yielded by the two approaches, were not correlated across the transplant sites. We argue that the ranking of site habitat suitability is probably more reliable in the transplant experiment than in the SDM because predictors in the former better describe understory conditions, but that ranking might vary among years, e.g. due to differences in climate. Our results suggest that L. vernus is limited by soil and light rather than directly by climate at its northern range edge, where conifers dominate forests and create suboptimal conditions of soil and canopy-penetrating light. A general implication of our study is that to better understand how climate change influences range dynamics, we should not only strive to improve existing approaches but also to use multiple approaches in concert.

Identifying the factors determining the abundance and distribution of species is a fundamental question in ecology. One key issue is how similar the factors determining species’ distributions across spatial scales are (here we focus especially on spatial extents). If the factors are similar across extents, then the large scale distribution pattern of a species may provide information about its local habitat requirements, and vice versa. We assessed the relationships between landscape and national optima as well as landscape and continental optima for growing degree days, maximum temperature and minimum temperature for 96 bryophytes and 50 vascular plants. For this set of species, we derived landscape optima from abundance weighted temperature data using species inventories in central Sweden and a fine-grained temperature model (50 m), national optima from niche centroid modelling based on GBIF data from Sweden and the same fine-grained climate model, and continental optima using the same method as for the national optima but from GBIF data from Europe and Worldclim temperatures (c. 1000 m). The landscape optima of all species were positively correlated with national as well as continental optima for maximum temperature (r = 0.45 and 0.46, respectively), weakly so for growing degree days (r = 0.30 and r = 0.28), but sometimes absent for minimum temperature (r = 0.26 and r = 0.04). The regression slopes of national or continental optima on local optima did not differ between vascular plants and bryophytes for GDD and Tmax. However, the relationship between the optima of Tmin differed between groups, being positive in vascular plants but absent in bryophytes. Our results suggest that positive correlations between optima at different spatial scales are present for some climatic variables but not for others. Moreover, our results for vascular plants and bryophytes suggest that correlations might differ between organism groups and depend on the ecology of the focal organisms. This implies that it is not possible to routinely up- or downscale distribution patterns based on environmental correlations, since drivers of distribution patterns might differ across spatial extents.

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.

Variation in selection among populations and years has important implications for evolutionary trajectories of populations. Yet, the agents of selection causing this variation have rarely been identified. Selection on the time of reproduction within a season in plants might differ both among populations and among years, and selection can be mediated by both mutualists and antagonists. We investigated if differences in the direction of phenotypic selection on flowering phenology among 20 populations of Gentiana pneumonanthe during 2 yr were related to the presence of the butterfly seed predator Phengaris alcon, and if butterfly incidence was associated with the abundance of the butterfly's second host, Myrmica ants. In plant populations without the butterfly, phenotypic selection favored earlier flowering. In populations where the butterfly was present, caterpillars preferentially attacked early-flowering individuals, shifting the direction of selection to favoring later flowering. Butterfly incidence in plant populations increased with ant abundance. Our results demonstrate that antagonistic interactions can shift the direction of selection on flowering phenology, and suggest that such shifts might be associated with differences in the community context.

An insect species that shows variation in host species association across its geographical range may do so either because of local adaptation in host plant preference of the insect, or through envi ...

Time lags in responses of organisms to deteriorating environmental conditions delay population declines and extinctions. We examined how local processes at the population level contribute to extinction debt, and how cycles of habitat deterioration and recovery may delay extinction. We carried out a demographic analysis of the fate of the grassland perennial Primula veris after the cessation of grassland management, where we used either a unidirectional succession model for forest habitat or a rotation model with a period of forest growth followed by a clear-cut and a new successional cycle. The simulations indicated that P. veris populations may have an extinction time of decades to centuries after a detrimental management change. A survey of the current incidence and abundance of P. veris in sites with different histories of afforestation confirmed the simulation results of low extinction rates. P. veris had reduced incidence and abundance only at sites with at least 100 years of forest cover. Time to extinction in simulations was dependent on the duration of the periods with favourable and unfavourable conditions after management cessation, and the population sizes and growth rates in these periods. Our results thus suggest that the ability of a species to survive is a complex function of disturbance regimes, rates of successional change, and the demographic response to environmental changes. Detailed demographic studies over entire successional cycles are therefore essential to identify the environmental conditions that enable long-term persistence and to design management for species experiencing extinction debts.

In the present scenario of climatic change, climatic refugia will be of paramount importance for species per- sistence. Topography can generate a considerable climatic heterogeneity over short distances, which is often disregarded in macroclimatic predictive models. Here we investigate the role of rocky habitats as microclimatic refugia by combining two different analyses: exploring a thermal mechanism whereby rocky habitats might serve as refugia, and examining if the biogeographic pattern shows a high abundance of relict, endemic and peripheral species. The thermal profile of two populations of relict and endemic plant species occurring in Pyrenean cliffs was investigated by infrared images and in situ temperature data-loggers. Despite occurring in crevices of a south oriented slope, Androsace cylindrica showed a narrower daily range of temperature than the surrounding matrix, thereby avoiding extreme high temperatures. Borderea chouardii, of tropical ancestors, also occurred in patches where temperatures were buffered during the growth season, experiencing lower mean temperatures than the surrounding matrix and nearby areas during the warmer part of the day, and similar temperatures during the colder. The rocky habitats of both species, therefore, reduced temperature ranges and exposition to extreme climatic events. Compared to other habitats, the rocky ones also harboured a very high fraction of both endemics and peripheral plant populations according to the largest vegetation dataset available in the Pyrenees (18,800 plant inventories and 400,000 records). Our results suggest an association between the habitats of relicts, en- demics and species at their distribution limit, driven by a stabilizing effect of rocky habitats on extreme tem- peratures. Given the important role of rocky habitats as hotspots of singular and unique plants, their char- acterization seems a sensible first step to identify potential refugia in the context of climate change.

Pollination in gymnosperms is usually accomplished by means of wind, but some groups are insect-pollinated. We show that wind and insect pollination occur in the morphologically uniform genus Ephed ...

Little is known about the extent to which observed phenological responses to changes in climate are the result of phenotypic plasticity or genetic changes. We also know little about how plasticity, in terms of thermal reaction norms, vary spatially. We investigated if the thermal reaction norms for flower development of five crucifer species (Brassicaceae) differed among three regions along a south–north latitudinal gradient in replicated experiments. The mean response (elevation) of thermal reaction norms of flowering differed among regions in all study species, while sensitivity of flower development to temperature (slope) differed in only one of the species. Differences in mean responses corresponded to cogradient patterns in some species, but countergradient patterns in other. This suggests that differences among regions were not solely the result of adaptation to differences in the length of the growing season, but that other factors, such as herbivory, play an important role. Differences in development rate within species were mainly explained by variation in early phases of bud formation in some species but by variation in later phases of bud formation in other species. The differences in latitudinal patterns of thermal reaction norms among species observed in this study are important, both to identify agents of selection and to predict short‐ and long‐term responses to a warming climate., We thank Lars Ericson for providing information about plant populations, Patrick Gullström for field work assistance, the strategic research Programme EkoKlim at Stockholm Univ. for funding (to JE, KG, CW), and Séneca Foundation for funding (postdoctoral grant 12337/PD/09 to JANC)

Optimal timing of reproduction within a season may be influenced by several abiotic and biotic factors. These factors sometimes affect different components of fitness, making assessments of net selection difficult. We used estimates of offspring fitness to examine how pre-dispersal seed predation influences selection on flowering schedule in an herb with a bimodal flowering pattern, Actaea spicata. Within individuals, seeds from flowers on early terminal inflorescences had a higher germination rate and produced larger seedlings than seeds from flowers on late basal inflorescences. Reproductive value, estimated using demographic integral projection models and accounting for size-dependent differences in future performance, was two times higher for intact seeds from early flowers than for seeds from late flowers. Fruits from late flowers were, however, much more likely to escape seed predation than fruits from early flowers. Reproductive values of early and late flowers balanced at a predation intensity of 63%. Across 15 natural populations, the strength of selection for allocation to late flowers was positively correlated with mean seed predation intensity. Our results suggest that the optimal shape of the flowering schedule, in terms of the allocation between early and late flowers, is determined by the trade-off between offspring number and quality, and that variation in antagonistic interactions among populations influences the balancing of this trade-off. At the same time they illustrate that phenotypic selection analyses that fail to account for differences in offspring fitness might be misleading.

Timing of plant development both determines the abiotic conditions that the plant experiences and strongly influences the intensity of interactions with other organisms. Plants and herbivores differ in their response to environmental cues, and spatial and temporal variation in environmental conditions might influence the synchrony between host plants and herbivores, and the intensity of their interactions. We investigated whether differences in first day of flowering among and within 21 populations of the polyploid herb Cardamine pratensis influenced the frequency of oviposition by the butterfly Anthocharis cardamines during four study years. The proportion of plants that became oviposited upon differed among populations, but these differences were not related to mean flowering phenology within the population in any of the four study years. Attack rates in the field were also not correlated with resistance to oviposition estimated under controlled conditions. Within populations, the frequency of butterfly attack was higher in early-flowering individuals in two of the four study years, while there was no significant relationship in the other 2 years. Larger plants were more likely to become oviposited upon in all 4 years. The effects of first flowering day and size on the frequency of butterfly attack did not differ among populations. The results suggest that differences in attack intensities among populations are driven mainly by differences in the environmental context of populations while mean differences in plant traits play a minor role. The fact that within populations timing of flowering influenced the frequency of herbivore attack only in some years and suggests that herbivore-mediated selection on plant phenology differs among years, possibly because plants and herbivores respond differently to environmental cues.

Environmental changes are expected to alter both the distribution and the abundance of organisms. A disproportionate amount of past work has focused on distribution only, either documenting historical range shifts or predicting future occurrence patterns. However, simultaneous predictions of abundance and distribution across landscapes would be far more useful. To critically assess which approaches represent advances towards the goal of joint predictions of abundance and distribution, we review recent work on changing distributions and on effects of environmental drivers on single populations. Several methods have been used to predict changing distributions. Some of these can be easily modified to also predict abundance, but others cannot. In parallel, demographers have developed a much better understanding of how changing abiotic and biotic drivers will influence growth rate and abundance in single populations. However, this demographic work has rarely taken a landscape perspective and has largely ignored the effects of intraspecific density. We advocate a synthetic approach in which population models accounting for both density dependence and effects of environmental drivers are used to make integrated predictions of equilibrium abundance and distribution across entire landscapes. Such predictions would constitute an important step forward in assessing the ecological consequences of environmental changes.

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.

Diapause plays a central role in insect life cycles by allowing survival during adverse seasonal conditions as well as synchronizing life cycles with the period of mate and food availability. Seasonal timing is expected to be particularly important for species that are dependent on resources available during a short time window-so-called phenological specialists-and latitudinal clines in seasonality are expected to favor local adaptation in phenological timing. However, to what degree latitudinal variation in diapause dynamics and post-winter development due to such local adaptation is influenced by the degree of phenological specialization is not well known. We experimentally studied two pierid butterfly species and found that the phenological specialist Anthocharis cardamines had shorter diapause duration than the phenological generalist Pieris napi along a latitudinal gradient in Sweden. Moreover, diapause duration increased with latitude in P. napi but not in A. cardamines. Sensitivity of the two species to winter thermal conditions also differed; additional cold temperature during the winter period shortened diapause duration for P. napi pupae but not for A. cardamines pupae. In both species, post-winter pupal development was faster after longer periods of cold conditions, and more southern populations developed faster than northern populations. Post-winter development was also invariably faster at higher temperatures in both species. We argue that the observed differences in diapause dynamics between the two species might be explained by the difference in phenological specialization that influences the costs of breaking diapause too early in the season.

Latitudinal clines in thermal reaction norms of development are a common phenomenon in temperate insects. Populations from higher latitudes often develop faster throughout the range of relevant tem ...

The main way in which plants can exert control over their local environment is by the timing of different events within their life cycles. Regarding timing of flowering as an integrated part of both the annual cycle and of the whole life cycle, rather than as an isolated event, has important implications for how we assess selection on timing of reproduction and interpret existing phenological patterns in perennial plants. I argue that: 1) we have little unequivocal evidence of pollinator-mediated selection on flowering time, but perhaps more evidence of antagonist-mediated selection; 2) much of selection on flowering time might occur before flowers have developed and after reproduction; 3) vital rates of non-flowering individuals can influence the strength and direction of selection on flowering time, and 4) differences in the direction of selection on flowering date between years might well correspond to consistent selection on the mechanisms determining flowering time. Overall, a life cycle perspective on timing of flowering is likely to facilitate the identification of selective agents and the understanding of the complex mechanisms underlying spatial and temporal variation in selection as well as to enable more accurate predictions of responses to environmental change.

Variation in mutualistic and antagonistic interactions are important sources of variation in population dynamics and natural selection. Environmental heterogeneity can influence the outcome of interactions by affecting the intensity of interactions, but also by affecting the demography of the populations involved. However, little is known about the relative importance of environmental effects on interaction intensities and demographic sensitivity for variation in population growth rates. We investigated how soil depth, soil moisture, soil nutrient composition, and vegetation height influenced the intensity of seed predation as well as host plant demography and sensitivity to seed predation in the perennial herb Primula farinosa. Intensity of seed predation ranged from 0% to 80% of seeds damaged among the 24 study populations and was related to soil moisture in two of four years. The effect of seed predation on plant population growth rate (lambda) ranged from negligible to a reduction in lambda by 0.70. Sensitivity of population growth rate to predation explained as much of the variation in the reductions in population growth rate due to seed predation as did predation intensity. Plant population growth rate in the absence of seed predation and sensitivity to predation were negatively related to soil depth and soil moisture. Both intensity of predation and sensitivity to predation were positively correlated with potential population growth rate and, as a result, there was no significant relationship between predation intensity and realized population growth rate. We conclude that in our study system environmental context influences the effects of seed predation on plant fitness and population dynamics in two important ways: through variation in interaction intensity and through sensitivity to the effects of this interaction. Moreover, our results show that a given abiotic factor can influence population growth rate in different directions through effects on potential growth rate, intensity of biotic interactions, and the sensitivity of population growth rate to interactions.

Patterns of sex expression and sex ratios are key features of the life histories of organisms. Bryophytes are the only haploid-dominant land plants. In contrast with seed plants, more than half of ...

Uncertainty associated with ecological forecasts has long been recognized, but forecast accuracy is rarely quantified. We evaluated how well data on 82 populations of 20 species of plants spanning ...

Accepted manuscript version. Published version available at https://doi.org/10.1016/j.ppees.2017.09.008. Accepted manuscript version, licensed CC BY-NC-ND 4.0. In the face of climate change, populations have two survival options − they can remain in situ and tolerate the new climatic conditions (“stay”), or they can move to track their climatic niches (“go”). For sessile and small-stature organisms like alpine plants, staying requires broad climatic tolerances, realized niche shifts due to changing biotic interactions, acclimation through plasticity, or rapid genetic adaptation. Going, in contrast, requires good dispersal and colonization capacities. Neither the magnitude of climate change experienced locally nor the capacities required for staying/going in response to climate change are constant across landscapes, and both aspects may be strongly affected by local microclimatic variation associated with topographic complexity. We combine ideas from population and community ecology to discuss the effects of topographic complexity in the landscape on the immediate “stay” or “go” opportunities of local populations and communities, and on the selective pressures that may have shaped the stay or go capacities of the species occupying contrasting landscapes. We demonstrate, using example landscapes of different topographical complexity, how species’ thermal niches could be distributed across these landscapes, and how these, in turn, may affect many population and community ecological processes that are related to adaptation or dispersal. Focusing on treeless alpine or Arctic landscapes, where temperature is expected to be a strong determinant, our theorethical framework leads to the hypothesis that populations and communities of topographically complex (rough and patchy) landscapes should be both more resistant and more resilient to climate change than those of topographically simple (flat and homogeneous) landscapes. Our theorethical framework further points to how meta-community dynamics such as mass effects in topographically complex landscapes and extinction lags in simple landscapes, may mask and delay the long-term outcomes of these landscape differences under rapidly changing climates.

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.

Mutualistic and antagonistic interactions with animals are known to influence the performance of plants in many ways. Much less is known about how such effects are influenced by the environment and how they translate into effects on plant population dynamics. In this study, we first quantified how pre-dispersal seed predation in the perennial herb Primula veris changes along a continuous gradient of canopy cover. We then used a deterministic demographic modeling approach to investigate how seed predation may influence population growth rate (λ) and how effects depend on environmentally-induced variation in plant demography. Intensity of seed predation increased with the degree of canopy cover, while sensitivity of λ to changes in seed production decreased. This translated into non-linear effects of seed predation on λ along the canopy cover gradient. Despite seed predation rates being highest in closed habitats, the negative effect of seed predation on λ was lower here than in slightly more open habitats. Our results demonstrate that knowledge of the intensity of plant-herbivore interactions does not suffice to infer how animals influence the population dynamics of plants and their distribution. Plant demographic sensitivity and its dependence on the environment need also to be taken into account when assessing the importance of plant-animal interactions. © Oikos.

Identifying the internal and external drivers of population dynamics is a key objective in ecology, currently accentuated by the need to forecast the effects of climate change on species distributions and abundances. The interplay between environmental and density effects is one particularly important aspect of such forecasts. We examined the simultaneous impact of climate and intraspecific density on vital rates of the dwarf shrub Fumana procumbens over 20 yr, using generalized additive mixed models. We then analyzed effects on population dynamics using integral projection models. The population projection models accurately captured observed fluctuations in population size. Our analyses suggested the population was intrinsically regulated but with annual fluctuations in response to variation in weather. Simulations showed that implicitly assuming variation in demographic rates to be driven solely by the environment can overestimate extinction risks if there is density dependence. We conclude that density regulation can dampen effects of climate change on Fumana population size, and discuss the need to quantify density dependence in predictions of population responses to environmental changes.

The relationship between the performance of individuals and the surrounding environment is fundamental in ecology and evolutionary biology. Assessing how abiotic and biotic environmental factors influence demographic processes is necessary to understand and predict population dynamics, as well as species distributions and abundances. We searched the literature for studies that have linked abiotic and biotic environmental factors to vital rates and, using structured demographic models, population growth rates of plants. We found 136 studies that had examined the environmental drivers of plant demography. The number of studies has been increasing rapidly in recent years. Based on the reviewed studies, we identify and discuss several major gaps in our knowledge of environmentally driven demography of plants. We argue that some drivers may have been underexplored and that the full potential of spatially and temporally replicated studies may not have been realized. We also stress the need to employ relevant statistical methods and experiments to correctly identify drivers. Moreover, assessments of the relationship between drivers and vital rates need to consider interactive, nonlinear and indirect effects, as well as effects of intraspecific density dependence. Synthesis. Much progress has already been made by using structured population models to link the performance of individuals to the surrounding environment. However, by improving the design and analyses of future studies, we can substantially increase our ability to predict changes in plant population dynamics, abundances and distributions in response to changes in specific environmental drivers. Future environmentally explicit demographic models should also address how genetic changes prompted by selection imposed by environmental changes will alter population trajectories in the face of continued environmental change and investigate the reciprocal feedback between plants and their biotic drivers.

The shape of the relationship between intensity of biotic interactions and strength of selection is important for spatial variation in selection, but is little explored. We quantified interactions and selection in 69 populations of the short-lived herb Primula farinosa. As predicted because of saturation and depletion effects, the strength of selection on a discrete and on a continuously varying floral display trait were in several cases significantly non-linearly related to the mean intensity of mutualistic and antagonistic interactions. Strength of selection was strongest at low levels of fruit initiation and at high intensities of seed predation. Seed predation varied more among populations than did fruit initiation and could explain a larger proportion of the among-population variation in strength of selection. Our results support the contention that interaction intensity affects selection strength, and suggests that for mutualistic and antagonistic interactions that can be saturated or depleted, this relationship is sometimes non-linear.

Many recent studies have explored the effects of present and past landscape structure on species distribution and diversity. However, we know little about the effects of past landscape structure on ...

Costs of reproduction should depend on resource availability and on reproductive effort, which in turn may depend on traits influencing reproductive success. Therefore, variation in both habitat qu ...