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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).

Context Both climatic extremes and land-use change constitute severe threats to biodiversity, but their interactive effects remain poorly understood. In forest ecosystems, the effects of climatic extremes can be exacerbated at forest edges. Objectives We explored the hypothesis that an extreme summer drought reduced the richness and coverage of old-growth forest species, particularly in forest patches with high edge exposure. Methods Using a high-resolution spatially explicit precipitation dataset, we could detect variability in drought intensity during the summer drought of 2018. We selected 60 old-growth boreal forest patches in central Sweden that differed in their level of drought intensity and amount of edge exposure. The year after the drought, we surveyed red-listed and old-growth forest indicator species of vascular plants, lichens and bryophytes. We assessed if species richness, composition, and coverage were related to drought intensity, edge exposure, and their interaction. Results Species richness was negatively related to drought intensity in forest patches with a high edge exposure, but not in patches with less edge exposure. Patterns differed among organism groups and were strongest for cyanolichens, epiphytes associated with high-pH bark, and species occurring on convex substrates such as trees and logs. Conclusions Our results show that the effects of an extreme climatic event on forest species can vary strongly across a landscape. Edge exposed old-growth forest patches are more at risk under extreme climatic events than those in continuous forests. This suggest that maintaining buffer zones around forest patches with high conservation values should be an important conservation measure.

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.

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.

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.

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.

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.

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 ...

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.

Knowledge of how species interactions are influenced by climate warming is paramount to understand current biodiversity changes. We review phenological changes of Swedish butterflies during the latest decades and explore potential climate effects on butterfly–host plant interactions using the Orange tip butterfly Anthocharis cardamines and its host plants as a model system. This butterfly has advanced its appearance dates substantially, and its mean flight date shows a positive correlation with latitude. We show that there is a large latitudinal variation in host use and that butterfly populations select plant individuals based on their flowering phenology. We conclude that A. cardamines is a phenological specialist but a host species generalist. This implies that thermal plasticity for spring development influences host utilization of the butterfly through effects on the phenological matching with its host plants. However, the host utilization strategy of A. cardamines appears to render it resilient to relatively large variation in climate. Electronic supplementary material The online version of this article (doi:10.1007/s13280-014-0602-z) contains supplementary material, which is available to authorized users.

Microrefugia are sites that support populations of species when their ranges contract during unfavorable climate episodes. Here, we review and discuss two aspects relevant for microrefugia. First, distributions of different species are influenced by different climatic variables. Second, climatic variables differ in the degree of local decoupling from the regional climate. Based on this, we suggest that only species limited by climatic conditions decoupled from the regional climate can benefit from microrefugia. We argue that this restriction has received little attention in spite of its importance for microrefugia as a mechanism for species resilience (the survival of unfavorable episodes and subsequent range expansion). Presence of microrefugia will depend on both the responses of individual species to local climatic variation and how climate-forcing factors shape the correlation between local and regional climate across space and time.

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 ...

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.

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 ...

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.

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 ...

This study examined the interactive effects of early and late season herbivory on the growth and reproductive output of figwort (Scrophularia nodosa). The early season herbivore is a pentatomid bug that feeds on and kills the apical meristem, while the late season herbivores are 2 weevil species and a sawfly that all feed on leaves and flowers. The direct effect of early season meristem damage on plant reproduction was quite limited, although meristem damage did cause increased branching. This change in plant morphology may entail that early season herbivores have profound indirect effects on plant reproduction by affecting the abundance of and damage caused by late season herbivores. Comparisons of plants with and without early season meristem damage, natural and artificial, also suggest that plants with meristem damage are significantly shorter throughout most of the summer and receive less damage late in season. However, the reduced damage translated to increased flowering but not to increased fruit production, suggesting that the plants were able to compensate for late season damage. In the end, and despite damage, figwort was well able to tolerate the observed meristem and leaf damage.

Summary 1. Understanding how vital rates and reproductive value change with age is fundamental to demography, life history evolution and population genetics. The universality of organism senescence has been questioned on both theoretical and empirical grounds, and the prevalence and strength of senescence remain a controversial issue. Plants are particularly interesting for studies of senescence since individuals of many species have been reported to reach very high ages. 2. In this study, we examined whether the herb Borderea pyrenaica, known to reach ages of more than 300 years, experiences senescence. We collected detailed demographic information from male and female individuals in two populations over 5 years. An unusual morphological feature in this species enabled us to obtain exact age estimates for each of the individuals at the end of the demographic study. 3. We used restricted cubic regression splines and generalized linear models to determine nonlinear effects of age and size on vital rates. We then incorporated the effects of age and size in integral projection models of demography for determining the relationship between age and reproductive value. As the species is dioecious, we performed analyses separately for males and females and examined also the hypothesis that a larger reproductive effort in females comes at a senescence cost. 4. We found no evidence for senescence. Recorded individuals reached 260 years, but growth and fecundity of female and male individuals did not decrease at high ages, and survival and reproductive value increased with age. The results were qualitatively similar also when accounting for size and among-individual vital rate heterogeneity, with the exception that male flowering probability decreased with age when accounting for size increases. 5.Synthesis. Overall, our results show that performance of both male and female plants of B. pyrenaica may increase rather than decrease at ages up to several centuries, and they support the notion that senescence may be negligible in long-lived modular organisms. This highlights the need to explore mechanisms that enable some species to maintain high reproductive values also at very high ages and to identify the evolutionary reasons why some organisms appear to experience no or negligible senescence.

7 páginas, 3 figuras, 1 tabla.-- El documento en word es la versión de autor., To accurately estimate population dynamics and viability, structured population models account for among-individual differences in demographic parameters that are related to individual state. In the widely used matrix models, such differences are incorporated in terms of discrete state categories, whereas integral projection models (IPMs) use continuous state variables to avoid artificial classes. In IPMs, and sometimes also in matrix models, parameterization is based on regressions that do not always model nonlinear relationships between demographic parameters and state variables. We stress the importance of testing for nonlinearity and propose using restricted cubic splines in order to allow for a wide variety of relationships in regressions and demographic models. For the plant Borderea pyrenaica, we found that vital rate relationships with size and age were nonlinear and that the parameterization method had large effects on predicted population growth rates, λ (linear IPM, 0.95; nonlinear IPMs, 1.00; matrix model, 0.96). Our results suggest that restricted cubic spline models are more reliable than linear or polynomial models. Because even weak nonlinearity in relationships between vital rates and state variables can have large effects on model predictions, we suggest that restricted cubic regression splines should be considered for parameterizing models of population dynamics whenever linearity cannot be assumed., This study was financially supported by the Strategic Research Programme EkoKlim at Stockholm University (J.P.D.), the National Parks project (ref 018/2008) (M.B.G), and the Swedish Research Council for Environment, Agricultural Sciences and Spatial Planning (FORMAS) (J.E.).

Despite seemingly obvious effects of environmental drivers, mechanisms behind long-term changes in plant population sizes over time are often poorly known. We investigated how soil potassium concentration and seed predation are likely to change over time as a result of succession from deciduous forest to spruce forest, and how this affects population trajectories of Actaea spicata. Observations and addition experiments showed that high soil potassium concentration increased individual growth rates. Among-site comparisons showed that soil potassium concentration was lower where proportion spruce was higher. Incorporation of a gradual increase in spruce over time in an integral projection model where individual growth depended on potassium suggested a net decrease in A. spicata population sizes over forest succession. This result suggests that small changes in factors with small effects on individual performance can influence patterns of species occupancy along successional gradients. We incorporated also density independent and density dependent effects of pre-dispersal seed predation over succession into the same model. Seed predation influenced the tree composition at which A. spicata population growth was positive. However, significant effects of A. spicata population size on seed predation intensity did not translate into important feedback effects on population growth trajectories over succession. Our results illustrate how demographic models can be used to gain understanding of the mechanisms behind effects of environmental change on species abundances and distributions by the simultaneous inclusion of changing abiotic and biotic factors.

Negative effects of habitat fragmentation on individual performance have been widely documented, but relatively little is known about how simultaneous effects on multiple vital rates translate into effects on population viability in long-lived species. In this study, we examined relationships between population size, individual growth, survival and reproduction, and population growth rate in the perennial plant Phyteuma spicatum. Population size positively affected the growth of seedlings, the survival of juveniles, the proportion of adults flowering, and potential seed production. Analyses with integral projection models, however, showed no relationship between population size and population growth rate. This was due to the fact that herbivores and pathogens eliminated the relationship between population size and seed production, and that population growth rate was not sensitive to changes in the vital rates that varied with population size. We conclude that effects of population size on vital rates must not translate into effects on population growth rate, and that populations of long-lived organisms may partly be able to buffer negative effects of small population size on vital rates that have a relatively small influence on population growth rate. Our study illustrates that we need to be cautious when assessing the consequences of habitat fragmentation for population viability based on effects on only one or a few vital rates.

Explaining variation in population growth rates is fundamental to predicting population dynamics and population responses to environmental change. In this study, we used matrix population models, which link birth, growth and survival to population growth rate, to examine how and why population growth rates vary within and among 50 terrestrial plant species. Population growth rates were more similar within species than among species; with phylogeny having a minimal influence on among-species variation. Most population growth rates decreased over the observation period and were negatively autocorrelated between years; that is, higher than average population growth rates tended to be followed by lower than average population growth rates. Population growth rates varied more through time than space; this temporal variation was due mostly to variation in post-seedling survival and for a subset of species was partly explained by response to environmental factors, such as fire and herbivory. Stochastic population growth rates departed from mean matrix population growth rate for temporally autocorrelated environments. Our findings indicate that demographic data and models of closely related plant species cannot necessarily be used to make recommendations for conservation or control, and that post-seedling survival and the sequence of environmental conditions are critical for determining plant population growth rate.

Summary 1. A primary goal of evolutionary ecology is to understand factors selecting for the diversity of life histories. Life-history components, such as time-to-reproduction, adult survivorship and fecundity, might differ among species because of variation in direct and indirect benefits of these life histories in different environments or might have lower-than-expected variability because of phylogenetic constraints. Here, we present a phylogenetic examination of demography and life histories using a data base of 204 terrestrial plant species. 2. Overall, statistical models without phylogeny were preferred to models with phylogeny for vital rates and elasticities, suggesting that they lacked phylogenetic signal and are evolutionarily labile. However, the effect of phylogeny was significant in models including sensitivities, suggesting that sensitivities exhibit greater phylogenetic signal than vital rates or elasticities. 3. Species with a greater age at first reproduction had lower fecundity, consistent with a cost of delayed reproduction, but only in some habitats (e.g. grassland). We found no evidence for an indirect benefit of delayed reproduction via a decrease in variation in fecundity with age to first reproduction. 4. The greater sensitivity and lower variation in survival than in fecundity was consistent with buffering of more important vital rates, as others have also found. This suggests that studies of life-history evolution should include survival, rather than only fecundity, for the majority of species. 5. Synthesis. Demographic matrix models can provide informative tests of life-history theory because of their shared construction and outputs and their widespread use among plant ecologists. Our comparative analysis suggested that there is a cost of delayed reproduction and that more important vital rates exhibit lower variability. The absolute importance of vital rates to population growth rates (sensitivities) exhibited phylogenetic signal, suggesting that a thorough understanding of life-history evolution might require an understanding of the importance of vital rates, not just their means, and the role of phylogenetic history.

Summary 1. In the understorey herb Lathyrus vernus seed production is pollen limited, but increased reproduction results in a lower probability of remaining reproductive. Putting these two results together, previous research reported that population growth rate λ was negatively impacted by high pollination. 2. Thus, costs and benefits have to be translated into the common currency of their respective effects on population dynamics to determine whether populations are truly pollen limited or whether they are already at an optimal level of pollination. 3. Also, when pollinators and demography vary from year to year we require a framework that examines reproductive benefits and demographic costs in the context of a variable environment. Whether or not additional pollination will increase the stochastic population growth rate λS depends upon the balance of stochastic elasticities of the costs and benefits. 4. In constant environment models, where seed survival, germination and seedling survival were increased, we found that the high cost of reproduction could be offset by improvements in seed survival and germination, but not by improvements of seedling survival. 5. In variable environment models, where changes in the sequence and frequencies of high- and low-pollination years mixed with occasional high-germination years were modelled, we found that increasing the frequency of high-germination conditions could offset the cost of reproduction, and the offset was even greater if high-germination years occurred after a high-pollination year or if high pollination was accompanied by high-germination conditions in the same year. 6. Both deterministic λ and stochastic λS were less sensitive to perturbation of reproduction than to perturbation of the probability for flowering plants to remain reproductive. In other words, a small change in the parameter which is related to the ‘cost’ of reproduction had a bigger effect than a small change in the parameter which is related to the ‘benefit’ of increased pollination for Lathyrus. 7.Synthesis. Stochastic environment-specific elasticities for reproduction and stasis of flowering plants differ in their response to environmental context. The cost–benefit relationships, the ultimate fitness consequences of supplemental pollen, are influenced by the frequency and sequence of years differing in pollen availability and recruitment conditions.

We tested whether differences in ploidy level and previous exposure to herbivory can affect plant tolerance to herbivory. We conducted a common garden experiment with 12 populations of two ploidy levels of the perennial herb Cardamine pratensis (five populations of tetraploid ssp. pratensis and seven populations of octoploid ssp. paludosa). Earlier studies have shown that attack rates by the main herbivore, the orange tip butterfly Anthocharis cardamines, are lower in populations of octoploids than in populations of tetraploids, and vary among populations. In the common garden experiment, a combination of natural and artificial damage significantly reduced seed and flower production. We measured tolerance based on four plant-performance metrics: survival, growth, seed production and clonal reproduction. For three of these measurements, tolerance of damage did not differ between ploidy levels. For clonal reproduction, the octoploids had a higher tolerance than the tetraploids, although they experience lower herbivore attack rates in natural populations. Populations from sites with high levels of herbivory had higher tolerance, measured by seed production, than populations with low levels of herbivory. We did not detect any significant costs of tolerance. We conclude that high intensity of herbivory has selected for high tolerance measured by seed production in C. pratensis.

Sweden. Fluctuations of resources in time and space will influence not only species abundance but also interactions among species. For plantconsumer interactions, the effects of resource variation have mostly been studied in systems with high resource variability. Systems with moderate variations are less studied, although ecological and evolutionary dynamics of plants and consumers are likely to be affected also by less extreme variability. The effects of variation in a particular resource should depend on consumer diet width. We examined how spatial and temporal variation in seed production in the perennial herb Lathyrus vernus influenced population dynamics and resource utilization in two beetle pre-dispersal seed predators with different host ranges over six years. The monophagous Apion opeticum occupied fewer patches and had lower densities than the oligophagous Bruchus atomarius. The proportion of seeds attacked increased with increases in seed production between years for both seed predators. A possible explanation for these patterns is that population dynamics of beetles are driven largely by local factors and that the same factors influence both beetle performance and seed production. In B. atomarius, patterns may also be influenced by a more pronounced preference for L. vernus in years with a high seed production in L. vernus .W e conclude that relatively modest variation in seed production may result in responses that differ in both direction and extent from those usually observed in systems with high variation in seed production.

Summary 1. Although necessary for understanding and predicting population dynamics, abiotic and biotic interactions have only rarely been coupled to demography and population dynamics. 2. We estimated effects of 11 environmental factors on survival, growth and fertility of the perennial herb Actaea spicata and incorporated significant factors into integral projection models to assess their effect on population dynamics. 3. Statistical models suggested that high soil potassium concentration increased individual growth and that seed predation and, to a lesser extent, canopy cover reduced seed production. 4. Demographic models showed that both soil potassium concentration and pre-dispersal seed predation could reverse population growth from positive to negative. The observed range of soil potassium concentration corresponded to growth rates ( λ ) between 0.96 and 1.07, at mean observed seed predation intensity. At observed mean potassium concentration, growth rate ranged from 0.99 to 1.02 over observed seed predation intensities. 5. Sensitivity of population growth rate to different vital rates strongly influenced the relative effects of the two factors. Elasticity analysis suggested that proportional changes in soil potassium concentration result in seven times larger effects on population growth rate than changes in seed predation. 6. Synthesis . We conclude that relatively weak associations between environmental factors and vital rates can have substantial long-term effects on population growth. Approaches based on detailed demographic models, that simultaneously assess abiotic and biotic effects on population growth rate, constitute important tools for establishing the links between the environment and dynamics of populations and communities.

There is growing evidence that genetic and ecological factors interact in determining population persistence. The demographic effects of inbreeding depression can largely depend on the ecological milieu. We used demographic data of the perennial herb Succisa pratensis from six populations in grazed and ungrazed sites with different soil moisture. We built an individual-based model assessing the demographic consequences of inbreeding depression in populations with different management and habitat. Today this plant has to cope with severe landscape fragmentation, deteriorating habitat conditions in terms of decreasing grazing intensity, and the effects of inbreeding depression. For each population we performed simulations testing two inbreeding depression hypotheses (partial dominance and overdominance) and three epistatic functions among loci. The results indicated stronger inbreeding depression effects for populations in unfavourable sites without grazing or in xeric habitats compared to populations in favourable mesic sites with grazing. Overall, we found stronger effects with overdominance, a result that emphasizes the importance of understanding the genetic mechanisms of inbreeding depression. Hence, management practices can interact with the genetic consequences of inbreeding depression in population dynamics, which may have important implications for plant population ecology and evolutionary dynamics of inbreeding depression, [GER] Es gibt zunehmend Hinweise darauf, dass genetische und okologische Faktoren bei der Bestimmung der Persistenz von Populationen zusammenwirken. Die demographischen Effekte von Inzuchtdepressionen hangen in großem Ausmaß von der okologischen Umwelt ab. Wir nutzten die demographischen Daten des perennierenden Krautes Succisa pratensis aus sechs Populationen, die entweder beweidet oder unbeweidet waren und unterschiedliche Bodenfeuchten zeigten. Wir erstellten ein individuenbasiertes Modell um die demographischen Konsequenzen der Inzuchtdepression in verschiedenen Populationen bei verschiedenen Bewirtschaftungen und Habitaten abzuschatzen. Heute muss diese Pflanze mit ernster Landschaftsfragmentierung, sich verschlechternden Habitatbedingungen in Form von abnehmender Beweidungsintensitat und den Auswirkungen der Inzuchtdepression zurechtkommen. Fur jede Population fuhrten wir Simulationen durch, um zwei Inzuchtdepressions-Hypothesen (teilweise und Uberdominanz) und drei epistatische Funktionen uber die Loci zu testen. Die Ergebnisse wiesen auf starkere Auswirkungen der Inzuchtdepression bei Populationen in ungunstigen Habitaten ohne Beweidung oder in xerischen Habitaten im Vergleich zu Populationen in den bevorzugten mesischen Habitaten mit Beweidung hin. Insgesamt fanden wir starkere Effekte bei Uberdominanz, ein Ergebnis, dass die Wichtigkeit betont, die genetischen Mechanismen der Inzuchtdepression zu verstehen. Die Bewirtschaftungsweise kann also mit den genetischen Konsequenzen der Inzuchtdepression in der Populationsdynamik interagieren. Dies kann wichtige Implikationen fur die Pflanzenpopu- lationsokologie und die evolutionare Dynamik der Inzuchtdepression haben

Theory predicts that, with conflicting selection pressures mediated by mutualists and antagonists, alternative reproductive strategies can be maintained by negative frequency-dependent selection if it results in rare-morph advantage. We combined field experiments and surveys of natural populations to determine whether selection on floral display is frequency dependent in the self-incompatible herb Primula farinosa, which is polymorphic for inflorescence height and occurs in a short-scaped and a long-scaped morph. Among short-scaped plants, both pollination success, quantified as initiation of fruits and seeds, and seed predation were positively correlated with the relative frequency of the long-scaped morph. The relative strength of these effects and the direction of the resulting frequency-dependent selection on scape morph varied among years and populations. The results suggest that both mutualists and antagonists may mediate frequency-dependent selection and that frequency dependence may vary from positive to negative with rare-morph advantage, depending on the relative strength of these interactions.

Vicia pisiformis L. is a perennial leguminous plant with a main distribution in broadleaved forest-steppes of eastern Europe. The species is classified as endangered (EN) according to the IUCN red- ...

1. In patchily distributed plant species, seed production is likely to be influenced both by local abiotic factors affecting plant size and conditions for fruit maturation, and by population characteristics affecting the intensity of interactions with mutualists and antagonists. However, the relative importance of these effects is poorly known. 2. We used multiple regression and path models to examine the importance of abiotic factors (sun exposure, soil depth) and population characteristics (size, density and connectivity) for variation in flower and fruit production and intensity of seed predation among 39 populations of the long-lived herb Vincetoxicum hirundinaria in three consecutive years. In addition, we manipulated water availability in a field experiment and recorded short-term and long-term effects on fruit output, and conducted a supplemental hand-pollination experiment. 3. Flower production varied little, while fruit initiation, fruit abortion and fruit predation varied considerably among years. Sun exposure and soil depth affected fruit production per plant indirectly and positively through their effects on flower number. Population density affected fruit production negatively through its effect on flower number. Both fruit initiation and the proportion of fruits attacked by the tephritid fly Euphranta connexa were related positively to population size. 4. The number of full-size fruits per plant was related positively to sun exposure and population size in two years each, and related negatively to population density in one year. However, because of seed predation, the number of intact mature fruits was related significantly to population characteristics in only one of three years. 5. The field experiments showed that both shortage of water and insufficient pollination may limit fruit set in V. hirundinaria. 6. Synthesis. These results demonstrate that the relative importance of local abiotic conditions and population characteristics may vary considerably along the chain of events from flower formation to intact fruit, and also among years. They further show that, at least in species with a naturally patchy distribution, connectivity may be relatively unimportant for variation in reproductive output compared to effects of habitat quality, population size and density.