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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.

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

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.

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.

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.

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.

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 ...

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 ...

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.

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.

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 ...

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.

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.

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.

Evidence is accumulating that significant amounts of substances can move internally in ectohydric bryophytes, which lack specialized structures for internal water movement. This potential for resource translocation is likely to affect the productivity in individual shoot sections. We tested whether annual growth intervals in the pleurocarpous moss Pseudocalliergon trifarium were performing as independent units by comparing the biomass of annual increments within individual shoots from 13 populations from northern and central Sweden. Pseudocalliergon trifarium has a distinct shoot architecture with discrete annual segments and is therefore well suited to study annual biomass allocation patterns. Mass of the most recent growth interval was directly correlated with segment mass of at least two preceding years. The strength of the relationship between current growth and four consecutively older annual increments decreased linearly with age of the annual segments, suggesting that correlations within shoots were the result of resource translocation rather than of among-year correlations in local environmental factors. Our data suggest that interconnected segments are physiologically integrated, and that size relationships between successive increments are relatively independent of environmental conditions. We conclude that annual growth intervals cannot be considered as independent assimilatory units. Internal resource transfer across annual segments may buffer variation in external availability of or internal demands for resources, and has implications for the understanding of growth patterns and reproductive investments in ectohydric bryophytes.

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.

Pre-dispersal seed predation may have important effects on population dynamics and trait evolution in plants. In this review, we first present a conceptual framework of the strength of pre-dispersal seed predation and its variation in space and time. We consider the interaction between plants and their seed predators to be “strong” when it affects plant population dynamics or causes changes in plant trait–fitness relationships, and “weak” when it has no such effects, and propose ways of how to adequately assess these effects. Second, we review the ecological literature between 1991 and 2005 to evaluate documented effects of pre-dispersal seed predation on plants and draw five major conclusions. (1) Pre-dispersal seed predation rates are usually low but sometimes high, and show a considerable variation in space and time. (2) Direct evidence suggests that pre-dispersal seed predation can have a significant effect on recruitment and plant population growth rate. Accumulating evidence of seed-limited recruitment suggests that such effects are common. (3) Pre-dispersal seed predation affects selection on several plant traits, such as flowering phenology and flower number, which are usually interpreted mainly in the context of plant–pollinator interactions. (4) The patterns of variation in the interactions between plants and pre-dispersal seed predators suggest that geographic selection mosaics may be common. (5) Although there are numerous studies estimating seed predation, there are still rather few studies that have aimed at examining the interaction explicitly in terms of effects on plant population dynamics and trait selection. From these we know that seed predators can have important, and often variable, effects on plant population dynamics and trait evolution. However, it still remains to assess how important they are across study systems and relative to other aspects of the plant's biotic and abiotic environment.

Vegetative phenology constrains the onset of flowering in the perennial herb Lathyrus vernus

Background and Aims Roughly half of the species of bryophytes have separate sexes (dioecious) and half are hermaphroditic (monoecious). This variation has major consequences for the ecology and evolution of the different species. In some sexually reproducing dioecious bryophytes, sex ratio has been shown to vary with environmental conditions. This study focuses on the dioecious wetland moss Drepanocladus trifarius, which rarely produces sexual branches or sporophytes and lacks apparent secondary sex characteristics, and examines whether genetic sexes exhibit different habitat preferences, i.e. whether sexual niche partitioning occurs. Methods A total of 277 shoots of D. trifarius were randomly sampled at 214 locations and 12 environmental factors were quantified at each site. Sex was assigned to the individual shoots collected in the natural environments, regardless of their reproductive status, using a specifically designed molecular marker associated with female sex. Key Results Male and female shoots did not differ in shoot biomass, the sexes were randomly distributed with respect to each other, and environmental conditions at male and female sampling locations did not differ. Collectively, this demonstrates a lack of sexual niche segregation. Adult genetic sex ratio was female-biased, with 2·8 females for every male individual. Conclusions The results show that although the sexes of D. trifarius did not differ with regard to annual growth, spatial distribution or habitat requirements, the genetic sex ratio was nevertheless significantly female-biased. This supports the notion that factors other than sex-related differences in reproductive costs and sexual dimorphism can also drive the evolution of biased sex ratios in plants.

1 Seed limitation of plant species distribution has received much attention in recent years. The empirical evidence of seed limitation is, however, still weak, and mostly correlative. Unambiguous evidence can only be provided by long-term transplantation experiments. 2 We used an 11-year experiment, in which seeds of six perennial forest herbs were sown into 43 patches that were either occupied or unoccupied by the target species, to investigate the importance of seed limitation. In addition, we determined whether short-term (1–4 years) and long-term (11 years) experimental assessments provided similar results. Given that long-term experiments are generally not possible, we examined whether environmental factors and vegetation composition can correctly identify suitable habitats. 3 Our results show that the distribution of the forest herbs investigated were limited by the availability of seeds, and that this pattern was qualitatively consistent, over time. However, the number of initially unoccupied patches with successful recruitment decreased with time. 4 The fact that a similar trend in recruitment success was found in sowing experiments in initially occupied patches suggests that the decrease in recruitment success over time cannot solely be attributed to habitat suitability. Environmental factors, such as soil nutrients and vegetation composition, were in some cases correlated with recruitment success, but the strength of these relationships was often weak. 5 Our results support the notion that seed availability is an important limitation of plant species abundance and distribution. The results also point to several difficulties related to the empirical assessment of seed limitation. We therefore suggest a protocol that should enhance the quality of future experiments.

A fundamental assumption in life-history theory is that reproduction is costly. Higher reproductive investment for fruits than for flowers may result in larger costs of reproduction in females than in males, which is often used to explain male-skewed sex ratios in unisexual seed plants. In contrast, bryophytes have predominantly female-biased sex ratios, suggested to be a product of a higher average cost of sexual reproduction in males. Empirical evidence to support this notion is largely lacking. We investigated sex-specific reproductive effort and costs in the unisexual moss Pseudocalliergon trifarium that has a female-dominated expressed sex ratio and rarely produces sporophytes. Annual vegetative segment mass did not differ among male, female, and non-expressing individuals, indicating that there was no threshold-size for sex expression. Mean and annual mass of sexual branches were higher in females than in males, but branch number per segment did not differ between sexes. Prefertilization reproductive effort for females was significantly greater (11.2%) than for males (8.6%). No cost for sexual branch production in terms of reduced relative vegetative growth or decreased investment in reproductive structures in consecutive years was detected. A higher realized reproductive cost in males cannot explain the unbalanced sex ratio in the study species.

Question: How accurately can a suite of suggested functional traits predict plant species response to succession from semiopen woodland to closed deciduous canopy forest? Location: Southeastern Sweden. Methods: Abundance of 46 field-layer plant species in a temperate deciduous forest, measured as frequency of occupied plots, was estimated in 1961, 1970 and 2003. Abundance change over time across species was tested for correlations with functional traits and literature information on habitat preference. Results: Increase in abundance was positively correlated with specific leaf area (SLA), weakly negatively correlated with seed mass and not significantly correlated with plant height or start, peak and length of the flowering period. Change in abundance was correlated with the Ellenberg light indicator value, whereas no correlations were found with Ellenberg values for nitrogen, calcium and moisture, or forest preference according to the literature. Conclusions: SLA was a better predictor of how field layer plants responded to succession from semi-open woodland to closed canopy forest than empirically-derived measures of habitat preference. The same holds for SLA in relation to seed size, indicating that interactions in the established life-cycle phase are more important than the recruitment phase for species response to succession.

Summary 1 We attempt to explain the current distribution of a long-lived perennial plant, Succisa pratensis , in a rural landscape in southern Sweden by linking its population biology with documented changes in the landscape, using a dynamical, spatially explicit model incorporating population dynamics and spatial spreading of the plant. Changes in the landscape were inferred from historical maps (1850 and 1900) and aerial photographs (1945 and 2001). 2 We tested whether predictions for the current species distribution are affected by assumptions about its early 19th century distribution, to determine whether recent history and current processes are dominant, and how past landscape changes determine current distributions. 3 Initial conditions influence predictions of current distribution, suggesting that the current distribution still partly reflects the distribution of the species in the early 19th century. A period of 150 years is too short for Succisa to have spread extensively if dispersal parameters are given realistic values. 4 Simulations in which present-day land-use patterns were imposed at earlier dates showed that changes in landscape structure over the past 175 years also had a strong effect on the present-day habitat occupancy and population sizes of Succisa . 5 The dominant process for Succisa now is extinction from marginal habitats. It is therefore likely that the (relatively) high present-day occupation patterns are still due to much larger areas having been available in the past rather than to successful dispersal. Although the species has responded to landscape changes, there is little evidence of population sizes reaching equilibrium. 6 Our approach shows that the wealth of landscape information available from historical maps can be linked with data on population biology by means of dynamical models that can make predictions about species dynamics.

Metapopulation theory (summarized by Hanski 1999) is one of the most influential developments in ecology of recent decades, for example as a cornerstone of conservation biology, and as a theoretical basis for studies of dispersal and of the distribution and abundance of species. A synthesis of metapopulation theory and landscape ecology has been regarded as a key to analyses of the ecological consequences of ongoing landscape changes (Wiens 1997). Although successfully applied to many regional systems of animal populations, the usefulness of metapopulation theory for studies of plants is still open to debate. As Husband & Barrett (1996) and Eriksson (1996) concluded, rather few studies have documented metapopulations in plants, and the same holds for other forms of regional dynamics, e.g. source-sink populations and remnant populations. In a recent paper, Freckleton & Watkinson (2002) have critically reviewed studies of plant metapopulations, with the objectives of assessing whether regional populations of plants really are metapopulations, and suggesting a new typology for different forms of plant regional dynamics. Freckleton & Watkinson stress, and we agree, that these issues are not merely semantic. The concepts of regional dynamics not only determine the ways in which empirical studies are performed, but also influence the interpretation of data (illustrated by Freckleton & Watkinson with studies claiming, but not showing, the existence of metapopulations) and the modelling of spatial dynamics of plant populations. Freckleton & Watkinson (2002) begin by listing four criteria that must be fulfilled in order to regard a regional population as a metapopulation: (i) suitable habitat occurs as discrete patches that may be occupied or unoccupied; (ii) all local populations have a measurable extinction risk; (iii) habitat patches are interconnected by dispersal, allowing the possibility of re-colonization; and (iv) local populations are not completely synchronized in their dynamics. If these criteria are met, regional dynamics cannot be inferred from local processes, and local dynamics cannot be understood without referring to regional patterns or processes. Freckleton & Watkinson (2002) conclude that, given these criteria, very few cases exist where plant metapopulations have been documented. Indeed, there are a number of reasons why plants may not develop metapopulations in the same way as short-lived mobile animals. The existence of long-lived life cycle stages (e.g. seeds or vegetative ramets) means that a local population may persist for a long time even though the patch has become unsuitable. Successful dispersal and recruitment may be very sporadic, and therefore recolonization is unlikely after local population extinction. Dispersal over long distances may be governed by chance events. The definition of 'long distance' may differ between species, but in many cases it may be no more than a few hundred metres (Cain et al. 2000). Suitable habitat is difficult to define, and suitability is likely to be a continuous function of abiotic and biotic environmental factors. Freckleton & Watkinson propose that, using a metapopulation concept for describing regional populations, many plant regional populations would be described as non-equilibrium metapopulations, a conclusion we also reached for plants in fragmented habitats (Eriksson & Ehrl~n 2001). We agree with Freckleton & Watkinson (2002) that it is difficult to assess whether plants in general fulfil the criteria for metapopulations. However, we disagree with their suggestions on a new typology for regional dynamics. We also believe that their conclusions concerning the difficulties in documenting metapopulations in plants may be interpreted as indicating that local processes are sufficient for understanding regional patterns of plant populations, although we acknowledge that this may not have been their intention. We have three objectives here: (i) to comment on the typology of regional dynamics suggested by Freckleton & Watkinson (2002); (ii) to suggest how regional processes might be of importance to plant distributions irrespective of whether plants have metapopulations; and (iii) to argue that metapopulation theory should be developed further as a tool for studies of plants, rather than being replaced by other concepts.

Summary 1 In a metapopulation context, the distribution of a species in a patchy landscape is interpreted as the equilibrium outcome of extinction and colonization processes. Populations are thus more likely to occur in larger, better connected habitat fragments. 2 To test whether metapopulation models explain distribution patterns of plant species with different life histories we investigated the incidence of herbaceous species in deciduous forests in four provinces of south Sweden. 3 These data were correlated with habitat quality (soil pH, nitrogen mineralization and organic matter) and habitat configuration (patch area and distance to the nearest deciduous forest patch). We also examined whether habitat configuration affected the distribution of species with different life history attributes. 4 All ground layer plant species at 81 sites were recorded, and distribution patterns of 57 species were tested against seed mass, seed number, presence of a seed bank, plant height, life span, mode of pollination, dispersal mode and habitat preference. 5 Habitat quality, especially pH, was more important for the incidence of species than habitat configuration with patch area and isolation significantly affecting only 11 and four species, respectively. Species favoured by larger area were also disadvantaged by greater isolation. 6 The importance of habitat configuration to a species varied with life history. Species that were more negatively affected by patch isolation tended to be habitat specialists and clonal perennials and to produce fewer seeds. Animal-dispersed species were more negatively affected by small stand size. 7 Habitat configuration may be less important for vascular plant distributions than habitat quality or the effects of land use history.

Most species distribution models assume a close link between climatic conditions and species distributions. Yet, we know little about the link between species’ geographical distributions and the sensitivity of performance to local environmental factors. We studied the performance of three bryophyte species transplanted at south- and north-facing slopes in a boreal forest landscape in Sweden. At the same sites, we measured both air and ground temperature. We hypothesized that the two southerly distributed species Eurhynchium angustirete and Herzogiella seligeri perform better on south-facing slopes and in warm conditions, and that the northerly distributed species Barbilophozia lycopodioides perform better on north-facing slopes and in relatively cool conditions. The northern, but not the two southern species, showed the predicted relationship with slope aspect. However, the performance of one of the two southern species was still enhanced by warm temperatures. An important reason for the inconsistent results can be that microclimatic gradients across landscapes are complex and influenced by many climate-forcing factors. Therefore, comparing only north- and south-facing slopes might not capture the complexity of microclimatic gradients. Population growth rates and potential distributions are the integrated results of all vital rates. Still, the study of selected vital rates constitutes an important first step to understand the relationship between population growth rates and geographical distributions and is essential to better predict how climate change influences species distributions. EkoKlim

Summary 1 A trade-off between current and future reproduction, often referred to as the cost of reproduction, is a fundamental assumption in life history theory. In long-lived plants, large absolute differences in size between individuals, storage of resources between reproductive events and organ preformation may make such costs difficult to demonstrate, especially when only natural variation is considered. 2 The long-lived legume Lathyrus vernus shows large size differences compared with variation in carbon resource allocation, and is known to store resources in below-ground rhizomes. We therefore followed individual plants over a period of 4 consecutive years. We examined the cost of reproductive investment by comparing the performance of untreated plants that differed in size and herbivore damage. We also compared controls with plants where we experimentally reduced flowering in terms of fitness measured as: survival, growth, flower number, fruit:flower ratio and storage. 3 Natural patterns of flowering and fruiting provided no evidence of a negative relationship between current and future reproduction. Individuals that produced fruits did not experience a lower probability of surviving and producing fruits the following season compared with flowering individuals that failed to produce any fruits, even when differences in above-ground size and herbivore damage were taken into account. 4 Flower removal in a single season increased the allocation to the rhizome but the size of shoot buds for the next season was not increased. Experimental manipulation of reproductive effort by repeated removal of flowers during 3 consecutive years, however, resulted in a significant increase in vegetative size and the probability of flowering and setting fruit compared with control plants. 5 While long-term data on natural variation in fruit production and short-term experimental data provided no evidence of a cost of reproduction, such a cost is still present, although detectable only after repeated flower removal.

We investigated the cost of sporophyte production in the moss Dicranum polysetum both by examining patterns of growth and reproduction in unmanipulated shoots and by experimentally manipulating sexual reproduction. The estimated proportion of total carbon investment allocated to sexual reproduction in sporophyte-producing shoots over the study period was 74.8%. Unmanipulated shoots that aborted all sporophytes had a significantly higher growth in the top shoots than shoots that produced sporophytes. In sporophyte-producing shoots, total apical growth decreased proportionately with the number of sporophytes. Experimental prevention of sporophyte development resulted in significantly higher total apical growth of the gametophytes. Shoots where current perichaetia were lacking when marked had a mass increase in the top shoots similar to manipulated sporophytic shoots whereas sporophytic control shoots grew significantly less than these two categories. The difference between control shoots and manipulated shoots in the mass of vegetative apical growth was mainly because of different length increments whereas mass per unit length was similar between groups. The probability to reproduce sexually in the year after the manipulation, and the biomass allocated to this reproduction, were not affected by the experimental treatment.

We experimentally investigated how pollinator- and herbivore-induced changes influence the performance of the long-lived herb Primula veris. Eight treatments that corresponded to natural factors normally affecting this species were designed to enhance or reduce reproductive success and resource availability (flower removal, supplementary pollination, defoliation). During the experimental season and in the following year we quantified responses in terms of survival, growth, and seed production of reproductive plants. Matrix population models were used to calculate population growth rate using the demographic parameters recorded in permanent plots and respective treatment groups. Seed production was not limited by pollen availability, and we found no evidence of a cost of reproduction. Leaf removal had either no effect or a negative effect on future performance, depending on the timing of removal. Defoliation early in the season reduced current seed production and future growth, whereas removal during fruit development affected performance in the following year. Demographic models suggest that leaf damage has a smaller negative impact than flower removal on overall performance in this population. Our results suggest that the source-sink paths vary over the season and that the timing of herbivory may influence the extent to which effects are carried over to subsequent reproductive seasons.

Timing of seasonal plant development can affect biotic interactions and plant fitness. Phenology is governed largely by temperature and may therefore be affected by global climate warming, making this an important area of research. Several factors in addition to temperature may cause differences in phenology. We studied the influence of local environment, plant size, and reproductive effort on shoot emergence and flowering time of 290 individuals of Actaea spicata (Ranunculaceae), distributed among 25 plots in four populations. We used multiple regression and structural equation models (SEM) to study causal relationships. Among plots, soil temperature and canopy cover explained 63% of the variation in shoot emergence. Soil temperature, slope, and canopy cover together explained 83% of the variation in flowering time. Within plots, small plants on steep south-facing slopes with high soil potassium concentrations emerged earlier in the year. Plants emerging earlier flowered earlier, but no environmental factors affected flowering time directly. We found no effects of reproductive effort. Our results support the view that flowering time of temperate forest herbs is constrained by several environmental factors acting indirectly through effects on shoot emergence time.

In plant populations where reproductive output is limited by pollinator visitation, plants with attractive floral displays should have a selective advantage. We examined the effect of inflorescence height on pollination success in Primula farinosa, which is dimorphic for scape length. To test the hypothesis that fruit and seed initiation are more strongly pollen-limited in the short-scaped than in the long-scaped morph, and that this difference is affected by spatio-temporal variation in pollen limitation, we conducted a hand-pollination experiment in four populations over 2 yr. Pollen limitation of fruit initiation varied among populations and years, and was stronger in the short-scaped than in the long-scaped morph. The results suggest that interactions with pollinators will need to be considered for a full understanding of the maintenance of this striking polymorphism. The study also shows that, although pollen limitation is likely to vary in space and time in many plant species, such variation is not necessarily associated with variation in selection on floral characters.

Plants have two principal defense mechanisms to decrease fitness losses to herbivory: tolerance, the ability to compensate fitness after damage, and resistance, the ability to avoid damage. Variation in intensity of herbivory among populations should result in variation in plant defense levels if tolerance and resistance are associated with costs. Yet little is known about how levels of tolerance are related to resistance and attack intensity in the field, and about the costs of tolerance. In this study, we used information about tolerance and resistance against larval herbivory by the butterfly Anthocharis cardamines under controlled conditions together with information about damage in the field for a large set of populations of the perennial plant Cardamine pratensis. Plant tolerance was estimated in a common garden experiment where plants were subjected to a combination of larval herbivory and clipping. We found no evidence of that the proportion of damage that was caused by larval feeding vs. clipping influenced plant responses. Damage treatments had a negative effect on the three measured fitness components and also resulted in an earlier flowering in the year after the attack. Tolerance was related to attack intensity in the population of origin, i.e. plants from populations with higher attack intensity were more likely to flower in the year following damage. However, we found no evidence of a relationship between tolerance and resistance. These results indicate that herbivory drives the evolution for increased tolerance, and that changes in tolerance are not linked to changes in resistance. We suggest that the simultaneous study of tolerance, attack intensity in the field and resistance constitutes a powerful tool to understand how plant strategies to avoid negative effects of herbivore damage evolve.

We investigated the effects of dispersal limitation, diaspore density, soil chemistry, disturbance and mollusc herbivory on the abundance and distribution of the perennial forest herb Dentaria bulbifera (Brassicaceae). After experimental sowing of bulbils in originally empty patches, juveniles of Dentaria emerged in 83% of plots. The seventh year after sowing, plants still persisted and were reproducing in several plots, demonstrating that the species is dispersal-limited. However, long-term survival of transplants was higher at sites where Dentaria occurred naturally, and this difference increased with time after sowing. Thus, the importance of local factors may only be expressed fully several years after colonisation. Differences in soil chemistry and disturbance treatment were not associated with differences in patch suitability. Removal of molluscs, however, significantly increased recruitment, and the effects of a single molluscicide treatment persisted after four years. We conclude that dis...

If we look at plant cover at a certain site, we can observe that it is made up of a number of species of different size and form, conferring a certain structure to the vegetation cover. If we compare two sites, we invariably see differences in structure and species composition. These differences are repeated across a hierarchy of scales: at the local scale, the regional scale and ultimately at the continental scale. By comparing structure and species compositions across a larger number of sites, a statistical pattern emerges from the data: some species are found together more frequently than expected and others seem to avoid each other. In fact, these positive and negative associations result in the observation that the species composition of a site is not just a random collection of species but is patterned, resulting in recognizable plant communities. In a spatial context we can fruitfully distinguish between within-site and among-site processes that influence the plant community. This distinction enables the description of species composition as a result of a dynamic process of within-community extinctions and colonizations from the outside. According to this view, community structure results from the differential performance of species as a function of biotic and abiotic environmental factors during the period between colonization and extinction. It now very much depends on the temporal scale of the dynamics of the system whether one puts emphasis on deterministic within-community processes, such as intraand interspecific competition, or on the stochastic processes of colonization and extinction. In fact there has been a long-standing debate about the relative importance of these two underlying assortative processes that are responsible for the perceived patterns in species composition (resulting in recognizable plant communities). This debate has its roots in the beginning of the last century and is conveniently summarized as the Clementsian versus the Gleasonian view on plant communities. The Clementsian view with its emphasis on the role of abiotic conditions and the regulative power of competitive processes has been dominant for a number of decades. In fact, a larger part of vegetation science is about finding the relationship between species composition and environmental factors. The fact that, given certain environmental conditions, we are able to

We investigated biomass allocation patterns in sporophyte-bearing and non-sporophytic plants of the perennial moss Dicranum polysetum Sw. from southern Sweden by comparing dry masses of various reproductive and vegetative plant modules. Assessing parental vegetative biomass is critical in bryophytes due to continuous decay of basal shoot portions. We propose a method to estimate the photosynthetically active gametophyte biomass at the onset of the current sporophyte cycle retrospectively. With this measure for vegetative plant size, the proportional investment in reproductive structures, reproductive effort, was 16% when sporophytes were produced, and 1.3% when perichaetia remained unfertilized. In sporophytic plants, vegetative apical growth and reproductive output (measured both as dry mass and sporophyte number) were positively correlated, and reproductive effort was inversely related to the mass of the annual segment preceding sporophyte initiation. The development of mature sporophytes was n...

Summary Seedling recruitment in the low-fecund perennial herb Lathyrus vernus was investigated during 4 years in 11 patches. Emergence and seedling survival were studied in unmanipulated quadrates and in quadrates where additional seeds had been sown. Experimental addition of seeds increased both the proportion of quadrates with recruitment and the number of seedlings within quadrates with recruitment. The probability of seedling emergence and seedling survival were not correlated with seed or seedling density. Emergence varied among patches but not among years. First-year survival of seedlings did not vary temporally or spatially. Seedling survival was correlated with initial seedling size, but not to germination phenology or herbivory. It is concluded that recruitment in L. vernus is seed limited and that reductions in seed production caused by seed predators and herbivores may influence the number of recruits and established individuals. However, factors other than seed availability also limit recruitment, and a concept of combined seed and microsite limitation of recruitment is therefore an appropriate interpretation of the observed patterns in L. vernus .