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Summary: Anthropogenetic climate change has caused range shifts among many species. Species distribution models (SDMs) are used to predict how species ranges may change in the future. However, most SDMs rarely consider how climate‐sensitive traits, such as phenology, which affect individuals' demography and fitness, may influence species' ranges.Using > 120 000 herbarium specimens representing 360 plant species distributed across the eastern United States, we developed a novel 'phenology‐informed' SDM that integrates phenological responses to changing climates. We compared the ranges of each species forecast by the phenology‐informed SDM with those from conventional SDMs. We further validated the modeling approach using hindcasting.When examining the range changes of all species, our phenology‐informed SDMs forecast less species loss and turnover under climate change than conventional SDMs. These results suggest that dynamic phenological responses of species may help them adjust their ecological niches and persist in their habitats as the climate changes.Plant phenology can modulate species' responses to climate change, mitigating its negative effects on species persistence. Further application of our framework will contribute to a generalized understanding of how traits affect species distributions along environmental gradients and facilitate the use of trait‐based SDMs across spatial and taxonomic scales. [ABSTRACT FROM AUTHOR]

Restricting seed collecting to local populations is a common practice in restoration because it is assumed that most plants are adapted to local environmental conditions. However, there is still considerable debate about whether local seed collection should be the default seed‐provenancing strategy as the effects of climate change are increasingly considered in restoration planning. It is especially important to explore whether local seed provenance is necessary for dominant species used in restoration because the success of these projects often rests on these species. Stipa pulchra is one such species that is commonly used in grassland restoration projects in California. To explore how different seed‐provenancing strategies affected the establishment and growth of S. pulchra, we established three common garden experiments distributed across a latitudinal gradient in California. We collected seeds from seven wild populations, germinated seeds in a common greenhouse, then planted all populations at the three common gardens. We assessed growth and reproduction for 2 years. We found limited evidence that restricting seed sourcing to local populations increased the establishment of S. pulchra compared to seeds from more distant populations. Instead, we found evidence to support the use of regional admixture seed sourcing to increase resiliency to environmental variation. In particular, we found being sourced from a dry location during the dry 2018 season was a benefit, highlighting the importance of including dry‐adapted populations to increase climate resilience. Our experiment highlights the importance of exploring multiple seed‐provenancing methods when designing a project to improve long‐term success. [ABSTRACT FROM AUTHOR]

Wild plant species provide excellent examples of qualitative traits that evolve in response to environmental challenges (e.g., flower color, heavy metal tolerance, cyanogenesis, and male sterility). In addition to such discrete characters, a dazzling array of continuously distributed, quantitative traits are expressed at every phase of the life cycle. These traits are known or suspected to have evolved by natural selection because they are heritable, differ among populations or closely related taxa occupying distinct habitats, and have individual phenotypes associated with survival and reproductive success. This special issue [American Journal of Botany 109(11)] focuses on the tools and approaches for detecting or inferring the ecological and genetic factors contributing to changes in genetically based variation of quantitative traits within or among populations, or causing their divergence among taxa. The assembled articles use one or more of three primary approaches to detect the process or outcome of natural selection on morphological, life history, reproductive, chemical, and physiological quantitative traits: the analysis of phenotypic or artificially imposed selection to detect direct and indirect selection on traits whose function is well‐understood; common garden experiments, including reciprocal transplants and "resurrection" experiments; and quantitative genetic analyses designed to detect and to estimate the environmental and genetic sources of phenotypic variation or to forecast short‐term evolutionary change. Together, these articles examine and reveal the adaptive capacity of quantitative traits and the genetically based constraints that may limit their directional evolutionary change, thereby informing and testing inferences, hypotheses, and predictions concerning the evolutionary trajectories of wild plant species. [ABSTRACT FROM AUTHOR]

Premise: In many species, seed size influences individual fitness, but its heritability is low, impeding its evolution. In heterogeneous environments, even if heritability of seed size is low, genetic variation in phenotypic plasticity for seed size may provide the opportunity for selection, but this possibility has rarely been investigated in wild species. The evolutionary trajectory of seed size depends on whether additive, maternal, or non‐additive genetic variance dominates; moreover, the expression of any of these sources of variance may be environment‐dependent, reflecting genetic variation in plasticity. In this study, we examined these sources of variation in seed size and their response to drought in Dithyrea californica. Methods: We used a diallel design to estimate variance components for seed size in three greenhouse‐raised populations sampled from California and northern Mexico. We replicated diallels in two watering treatments to examine genetic parameters and genotype × environment interactions affecting seed size. We estimated general (GCA) and specific (SCA) combining ability, reciprocal effects (RGCA and RSCA), and their interactions with water availability, and we sought evidence that sexual conflict influences seed size. Results: Norms of reaction revealed genetic variation in plasticity for seed size in each population. Seed size in D. californica is determined by the combination of watering treatment, GCA and RGCA; parental identity and water availability do not consistently affect seed size, and we detected no evidence for sexual conflict. Conclusions: Multiple sources of genetic variation in phenotypic plasticity for seed size have the potential to influence its evolutionary trajectory in heterogenous environments. [ABSTRACT FROM AUTHOR]

Premise: The study of phenotypic divergence of, and selection on, functional traits in closely related taxa provides the opportunity to detect the role of natural selection in driving diversification. If the strength or direction of selection in field populations differs between taxa in a pattern that is consistent with the phenotypic difference between them, then natural selection reinforces the divergence. Few studies have sought evidence for such concordance for physiological traits. Methods: Herbarium specimen records were used to detect phenological differences between sister taxa independent of the effects on flowering time of long‐term variation in the climate across collection sites. In the field, physiological divergence in photosynthetic rate, transpiration rate, and instantaneous water‐use efficiency were recorded during vegetative growth and flowering in 13 field populations of two taxon pairs of Clarkia, each comprising a self‐pollinating and a outcrossing taxon. Results: Historically, each selfing taxon flowered earlier than its outcrossing sister taxon, independent of the effects of local long‐term climatic conditions. Sister taxa differed in all focal traits, but the degree and (in one case) the direction of divergence depended on life stage. In general, self‐pollinating taxa had higher gas exchange rates, consistent with their earlier maturation. In 6 of 18 comparisons, patterns of selection were concordant with the phenotypic divergence (or lack thereof) between sister taxa. Conclusions: Patterns of selection on physiological traits measured in heterogeneous conditions do not reliably reflect divergence between sister taxa, underscoring the need for replicated studies of the direction of selection within and among taxa. [ABSTRACT FROM AUTHOR]

Herbarium records provide a broad spatial and temporal range with which to investigate plant responses to environmental change. Research on plant phenology and its sensitivity to climate has advanced with the increasing availability of digitized herbarium specimens, but limitations of specimen‐derived data can undermine the inferences derived from such research. One issue that has received little attention is collection site uncertainty (i.e. error distance), a measure of confidence in the location from which a specimen was collected. We conducted comparative analyses of phenoclimatic models to determine whether spatial deviations of 2, 5, 15 or 25 km between recorded and simulated collection sites, as well as the error distance reported in digitized records, affect estimates of the phenological sensitivity of flowering time to annual temperature and precipitation in a widespread annual California wildflower. In this approach, we considered both spatial and interannual variation in climatic conditions. Simulated site displacements led to increasingly weak estimates of phenological sensitivity to temperature and precipitation anomalies with increasing distances. However, we found no significant effect of reported error distance magnitude on estimates of phenological sensitivity to climate normals or anomalies. These findings suggest that the spatial uncertainty of collection sites among specimens of widely collected plant species may not adversely affect estimates of phenological sensitivity to climate, even though real discrepancies and georeferencing inaccuracy can negatively impact such estimates. Collection site uncertainty merits further attention as a potential source of noise in herbarium data, especially for research on how plant traits respond to spatial and interannual climatic variation. [ABSTRACT FROM AUTHOR]

Understanding the effects of climate change on the phenological structure of plant communities will require measuring variation in sensitivity among thousands of co‐occurring species across regions. Herbarium collections provide vast resources with which to do this, but may also exhibit biases as sources of phenological data. Despite general recognition of these caveats, validation of herbarium‐based estimates of phenological sensitivity against estimates obtained using field observations remains rare and limited in scope. Here, we leveraged extensive datasets of herbarium specimens and of field observations from the USA National Phenology Network for 21 species in the United States and, for each species, compared herbarium‐ and field‐based estimates of peak flowering dates expected under standardized temperature conditions, and of sensitivity of peak flowering time to geographic and interannual variation in mean minimum temperatures (TMIN). We found strong agreement between herbarium‐ and field‐based estimates for standardized peak flowering time (r = 0.91, p < 0.001) and for the direction and magnitude of sensitivity to both geographic TMIN variation (r = 0.88, p < 0.001) and interannual TMIN variation (r = 0.82, p < 0.001). This agreement was robust to substantial differences between datasets in 1) the long‐term TMIN conditions observed among collection and phenological monitoring sites and 2) the interannual TMIN conditions observed in the time periods encompassed by both datasets for most species. Our results show that herbarium‐based sensitivity estimates are reliable among species spanning a wide diversity of life histories and biomes, demonstrating their utility in a broad range of ecological contexts, and underscoring the potential of herbarium collections to enable phenoclimatic analysis at taxonomic and spatiotemporal scales not yet captured by observational data. [ABSTRACT FROM AUTHOR]

Offspring size is a key functional trait that can affect subsequent life history stages; in many species, it exhibits both local adaptation and phenotypic plasticity. Variation among populations in offspring size may be explained by various factors, including local climatic conditions. However, geographic variation in climate may be partitioned into long‐term and interannual sources of variation, which may differ in their effects on population mean offspring size. To assess environmental correlates of offspring size, we evaluated geographic variation in seed mass among 88 populations representing 6 species of Streptanthus (Brassicaceae) distributed across a broad climatic gradient in California. We examined the effects of temperature‐mediated growing season length and precipitation on population mean seed mass to determine whether it is best explained by (1) long‐term mean climatic conditions; (2) interannual climate anomalies (i.e., deviations in climate from long‐term means) during the year of seed development, or (3) interactions between climate variables. Both long‐term mean climate and climate anomalies in the year of collection were associated with population mean seed mass, but their effects differed in direction and magnitude. Relatively large seeds were produced at chronically wet sites but also during drier‐than‐average years. This contrast indicates that these associations may be generated by different mechanisms (i.e., adaptive evolution vs. phenotypic plasticity) and may be evidence of countergradient plasticity in seed mass. In addition, populations occurring in locations characterized by relatively long growing seasons produced comparatively large seeds, particularly among chronically dry sites. This study highlights the need to consider that the responses of seed mass to long‐term versus recent climatic conditions may differ and that climate variables may interact to predict seed mass. Such considerations are especially important when using these patterns to forecast the long‐ and short‐term responses of seed mass to climate change. The results presented here also contribute to our broader understanding of how climate drives long‐term (e.g., local adaptation) and short‐term (e.g., phenotypic plasticity) variation in functional traits, such as offspring size across landscapes. [ABSTRACT FROM AUTHOR]

Premise: Forecasting how species will respond phenologically to future changes in climate is a major challenge. Many studies have focused on estimating species‐ and community‐wide phenological sensitivities to climate to make such predictions, but sensitivities may vary within species, which could result in divergent phenological responses to climate change. Methods: We used 743 herbarium specimens of the mountain jewelflower (Streptanthus tortuosus, Brassicaceae) collected over 112 years to investigate whether individuals sampled from relatively warm vs. cool regions differ in their sensitivity to climate and whether this difference has resulted in divergent phenological shifts in response to climate warming. Results: During the past century, individuals sampled from warm regions exhibited a 20‐day advancement in flowering date; individuals in cool regions showed no evidence of a shift. We evaluated two potential drivers of these divergent responses: differences between regions in (1) the degree of phenological sensitivity to climate and (2) the magnitude of climate change experienced by plants, or (3) both. Plants sampled from warm regions were more sensitive to temperature‐related variables and were subjected to a greater degree of climate warming than those from cool regions; thus our results suggest that the greater temporal shift in flowering date in warm regions is driven by both of these factors. Conclusions: Our results are among the first to demonstrate that species exhibited intraspecific variation in sensitivity to climate and that this variation can contribute to divergent responses to climate change. Future studies attempting to forecast temporal shifts in phenology should consider intraspecific variation. [ABSTRACT FROM AUTHOR]

Adaptation to current and future climates can be constrained by trade‐offs between fitness‐related traits. Early seedling emergence often enhances plant fitness in seasonal environments, but if earlier emergence in response to seasonal cues is genetically correlated with lower potential to spread emergence among years (i.e., bet‐hedging), then this functional trade‐off could constrain adaptive evolution. Consequently, selection favoring both earlier within‐year emergence and greater spread of emergence among years—as is expected in more arid environments—may constrain adaptive responses to trait value combinations at which a performance gain in either function (i.e., evolving earlier within‐ or greater among‐year emergence) generates a performance loss in the other. All such trait value combinations that cannot be improved for both functions simultaneously are described as Pareto optimal and together constitute the Pareto front. To investigate how this potential emergence timing trade‐off might constrain adaptation to increasing aridity, we sourced seeds of two grasses, Stipa pulchra and Bromus diandrus, from multiple maternal lines within populations across an aridity gradient in California and examined their performance in a greenhouse experiment. We monitored emergence and assayed ungerminated seeds for viability to determine seed persistence, a metric of potential among‐year emergence spread. In both species, maternal lines with larger fractions of persistent seeds emerged later, indicating a trade‐off between within‐year emergence speed and potential among‐year emergence spread. In both species, populations on the Pareto front for both earlier emergence and larger seed persistence fraction occupied significantly more arid sites than populations off the Pareto front, consistent with the hypothesis that more arid sites impose the strongest selection for earlier within‐year emergence and greater among‐year emergence spread. Our results provide an example of how evaluating genetically based correlations within populations and applying Pareto optimality among populations can be used to detect evolutionary constraints and adaptation across environmental gradients. [ABSTRACT FROM AUTHOR]

In recent decades, the final frost dates of winter have advanced throughout North America, and many angiosperm taxa have simultaneously advanced their flowering times as the climate has warmed. Phenological advancement may reduce plant fitness, as flowering prior to the final frost date of the winter/spring transition may damage flower buds or open flowers, limiting fruit and seed production. The risk of floral exposure to frost in the recent past and in the future, however, also depends on whether the last day of winter frost is advancing more rapidly, or less rapidly, than the date of onset of flowering in response to climate warming. This study presents the first continental‐scale assessment of recent changes in frost risk to floral tissues, using digital records of 475,694 herbarium specimens representing 1,653 angiosperm species collected across North America from 1920 to 2015. For most species, among sites from which they have been collected, dates of last frost have advanced much more rapidly than flowering dates. As a result, frost risk has declined in 66% of sampled species. Moreover, exotic species consistently exhibit lower frost risk than native species, primarily because the former occupy warmer habitats where the annual frost‐free period begins earlier. While reducing the probability of exposure to frost has clear benefits for the survival of flower buds and flowers, such phenological advancement may disrupt other ecological processes across North America, including pollination, herbivory, and disease transmission. [ABSTRACT FROM AUTHOR]

The evolution of seed size may be influenced by intrinsic attributes of populations, such as mating system and extrinsic factors, such as climate. Several hypotheses propose that the evolution of self‐fertilization from an outcrossing progenitor will be accompanied by a reduction in seed size, but this prediction has not been rigorously tested. Many studies report that the mean seed size of populations or taxa is associated with long‐term climate conditions. Here, we examined the effects on seed size of both mating system and climate within a single genus.In the California wildflower genus, Clarkia (Onagraceae), we sampled seeds from 58 populations representing three pairs of sister taxa; each pair included a predominantly outcrossing and a facultatively selfing taxon. We then examined the independent effects on population mean seed size of mating system, elevation, long‐term (30‐year) climate conditions, and climate anomalies (the deviation between conditions in the year of collection and the long‐term mean), focusing on maximum monthly temperature (Tmax), cumulative moisture deficit and cumulative precipitation (PPT) during Clarkia's growing season (fall, winter and spring).In each taxon pair, the selfing taxon had smaller seeds than the outcrosser. Local, long‐term (1921–1980 and 1981–2000) mean Tmax, PPT and elevation were independently and negatively associated with seed size. Long‐term means for Tmax and PPT explain geographical variation in seed size better than climate anomalies in the year of collection.Synthesis. We corroborated two key hypotheses concerning the drivers of geographical variation in mean seed size. Small seeds in Clarkia co‐evolve with selfing (although the mechanism remains elusive) and in response to chronically warm and wet conditions. The effect of long‐term mean precipitation on seed size differs qualitatively from the effect of precipitation anomalies; relatively large seeds are produced in populations experiencing wetter‐than‐normal years. Ongoing climate change may therefore generate conflicting selection on seed size in Clarkia: intensifying drought is likely to lead to an evolutionary increase in seed size due to its effects on seedling survivorship, while climate‐driven declines in pollinators or selection favouring more rapid reproduction may promote the evolution of self‐pollination, facilitating the evolution of smaller seeds. [ABSTRACT FROM AUTHOR]

Premise: Herbarium specimens represent an outstanding source of material with which to study plant phenological changes in response to climate change. The fine‐scale phenological annotation of such specimens is nevertheless highly time consuming and requires substantial human investment and expertise, which are difficult to rapidly mobilize. Methods: We trained and evaluated new deep learning models to automate the detection, segmentation, and classification of four reproductive structures of Streptanthus tortuosus (flower buds, flowers, immature fruits, and mature fruits). We used a training data set of 21 digitized herbarium sheets for which the position and outlines of 1036 reproductive structures were annotated manually. We adjusted the hyperparameters of a mask R‐CNN (regional convolutional neural network) to this specific task and evaluated the resulting trained models for their ability to count reproductive structures and estimate their size. Results: The main outcome of our study is that the performance of detection and segmentation can vary significantly with: (i) the type of annotations used for training, (ii) the type of reproductive structures, and (iii) the size of the reproductive structures. In the case of Streptanthus tortuosus, the method can provide quite accurate estimates (77.9% of cases) of the number of reproductive structures, which is better estimated for flowers than for immature fruits and buds. The size estimation results are also encouraging, showing a difference of only a few millimeters between the predicted and actual sizes of buds and flowers. Discussion: This method has great potential for automating the analysis of reproductive structures in high‐resolution images of herbarium sheets. Deeper investigations regarding the taxonomic scalability of this approach and its potential improvement will be conducted in future work. [ABSTRACT FROM AUTHOR]

●Many angiosperms are hermaphroditic and produce bisexual flowers in which male (pollen export) and female (stigma receptivity) functions are separated temporally. This sequential hermaphroditism may be associated with variation in flower size, color, or pattern, all of which may influence pollinator attraction. In this study, we describe variation in these traits across discrete functional sex stages within and between 225 greenhouse‐grown individuals of Clarkia unguiculata (Onagraceae). In addition, to identify the effects of floral phenotype on pollinator attraction in this species, we examine the effects of these floral traits on pollen receipt in ~180 individuals in an experimental field array.●Petal area, ultraviolet (UV)‐absorbing nectar guide area, and blue and green mean petal reflectance differ significantly across the functional sex stages of C. unguiculata. Male‐ and female‐phase flowers display significantly different pollinator attraction traits. Petal and UV nectar guide area increase as flowers progress from male phase to female phase, while blue reflectance and green reflectance peak during anther maturation.●In field arrays of C. unguiculata, female‐phase flowers with large UV nectar guides receive more pollen than those with small nectar guides, and female‐phase flowers with high mean blue reflectance values are more likely to receive pollen than those with low blue reflectance. Female‐phase flowers with green mean reflectance values that differ most from background foliage also receive more pollen than those that are more similar to foliage. These findings indicate that components of flower color and pattern influence pollen receipt, independent of other plant attributes that may covary with floral traits. We discuss these results in the context of hypotheses that have been proposed to explain sex‐specific floral attraction traits, and we suggest future research that could improve our understanding of sexual dimorphism in sequentially hermaphroditic species and the evolution of features that promote outcrossing. [ABSTRACT FROM AUTHOR]

Aim: Climate affects the flowering time of many species. Little is known, however, about how climate influences the properties of regional floras, including the rate at which taxa flower sequentially throughout the flowering season. This study is the first to detect geographical variation in this rate across North America. In addition, we tested for the independent effects of intraspecific variation in flowering time and taxonomic composition on the rate of sequential flowering among regional floras distributed across a temperature gradient. Location: North America. Time period: This study examined >59,000 herbarium specimens that were collected in flower from 1901 to 2013. Major taxa studied: 2,803 angiosperm taxa. Methods: We identified 51 climatically homogeneous regions across the continental, mostly western states of the USA, in each of which ≥100 species were represented by herbarium specimens. We then examined the effects of mean annual temperature (MAT) on the rate of sequential flowering among species in each region. We also evaluated whether geographical variation in the rate of sequential flowering was attributable to intraspecific variation in the flowering time and/or the taxonomic composition of regional floras. Results: As MAT increased over space, the rate of sequential flowering (standardized by the absolute length of the flowering season in each region) increased among relatively early‐flowering taxa but decreased among the latest‐flowering taxa. Both intraspecific variation and shifts in taxonomic composition among floras contributed to this pattern. Main conclusions: Among floras throughout North America, the rate of sequential flowering among co‐occurring taxa changes with MAT. Intraspecific phenological variation primarily affects the rate of sequential flowering during the first half of the growing season, consistent with the inference that future warming will most strongly affect flowering synchrony among early‐flowering taxa. [ABSTRACT FROM AUTHOR]

Questions: All else being equal, populations of dioecious species with a 50:50 sex ratio have only half the effective reproductive population size of bisexual species of equal abundance. Consequently, there is a need to explain how dioecious and bisexual species coexist. Increased mean individual seed mass, fecundity, and population density have all been proposed as attributes of unisexual individuals or populations that may contribute to the persistence or resilience of dioecious species. To date, no studies have compared sympatric dioecious and cosexual species with respect to all three components of fitness. In this study, we sought evidence for these compensatory advantages (higher seed mass, greater seed production per unit basal area, and higher population density) in dioecious species. Location: Five 20–25 ha forest dynamic plots spanning a latitudinal gradient in China, including two temperate, two subtropical, and one tropical forest. Methods: We used a phylogenetically corrected generalized linear modelling approach to assess the phylogenetic dependence and joint evolution of sexual system, seed mass and production, and ecological abundances among 48–333 species and 32,568–136,237 individuals per forest. Results: Across all five forests, we detected no consistent advantage for dioecious relative to sympatric cosexual species with respect to mean individual seed mass, seed production or the density of stems in any size class. Conclusions: Our study suggests that seed traits may provide compensatory mechanisms in some forests, but most often the coexistence of sexual systems cannot be explained by advantages of dioecy related to seed quality and demographic parameters. Future investigations of the factors that promote coexistence may increase our understanding by expanding the search to include attributes such as lifespan and tolerance or resistance to herbivores. [ABSTRACT FROM AUTHOR]

Premise: Strong correlations between traits can obscure their independent effects on components of reproduction. Style length (SL) and petal area (PA) vary within species, for example, but their independent effects on the opportunity for selection among pollen genotypes are poorly understood. Previous work in Clarkia detected a positive effect of SL on pollen receipt, potentially intensifying selection. However, this apparent effect of SL may be influenced by a correlated trait, such as PA. Here, we examine the independent effects of these two traits on pollen receipt and performance. Methods: We collected petals and styles from wild populations of two insect‐pollinated Clarkia taxa and estimated the independent and combined effects of SL and PA on pollen receipt and performance. Results: In both taxa, SL and PA are positively correlated. In C. unguiculata, both traits positively and independently affect pollen receipt, but in C. xantiana ssp. xantiana, the two traits act only in combination to affect pollen receipt. In both taxa, pollen receipt positively affects the numbers of pollen tubes entering and penetrating the style, as well as pollen tube attrition. Conclusions: The effects of SL and PA on pollen receipt and performance are taxon specific. In C. unguiculata, both traits may be independent targets of selection due to their effects on pollen receipt. In C. xantiana ssp. xantiana, by contrast, the combined (but not independent) effects of SL and PA influence pollen receipt. Ecological differences between these taxa require exploration to understand the mechanisms by which these traits affect pollinator behavior. [ABSTRACT FROM AUTHOR]

Premise of the Study: Wild plant species that require the services of pollen‐feeding insects for reliable pollination may evolve features that attract and reward their mutualistic partners. Heterantherous species have been proposed to exhibit a "division of labor" whereby "feeding anthers" (which produce pollen that may be consumed by an insect) are distinguished from "reproductive anthers" (which produce pollen more likely to contribute to reproduction). In some heterantherous species, including Clarkia unguiculata (Onagraceae), these two anther types differ with respect to stamen length, anther size, pollen production, and pollen color. Methods: The primary goal of this study was to test one component of the "division of labor" hypothesis by comparing the performance of the pollen produced by each type of anther in C. unguiculata. To achieve this goal, under greenhouse conditions, we hand pollinated and assessed pollen performance (using epifluorescence microscopy) within ~228 flowers. Key Results: The pollen produced by the two anther types differed significantly with respect to both stigma and style penetration. The inner series of anthers produce pollen with higher performance than the outer series of longer, dark red anthers. Conclusions: These findings contradict previous descriptions of the genus, reporting that the inner diminutive series of anthers in Clarkia produce "abortive and nonfunctional" pollen. We outline the future research required to demonstrate the ecological function of heteranthery in this iconic wildflower group. [ABSTRACT FROM AUTHOR]

Determining the manner in which plant species shift their flowering times in response to climatic conditions is essential to understanding and forecasting the impacts of climate change on the world's flora. The limited taxonomic diversity and duration of most phenological datasets, however, have impeded a comprehensive, systematic determination of the best predictors of flowering phenology. Additionally, many studies of the relationship between climate conditions and plant phenology have included only a limited set of climate parameters that are often chosen a priori and may therefore overlook those parameters to which plants are most phenologically sensitive. This study harnesses 894,392 digital herbarium records and 1,959 in situ observations to produce the first assessment of the effects of a large number (25) of climate parameters on the flowering time of a very large number (2,468) of angiosperm taxa throughout North America. In addition, we compare the predictive capacity of phenological models constructed from the collection dates of herbarium specimens vs. repeated in situ observations of individual plants using a regression approach—elastic net regularization—that has not previously been used in phenological modeling, but exhibits several advantages over ordinary least squares and stepwise regression. When herbarium‐derived data and in situ phenological observations were used to predict flowering onset, the multivariate models based on each of these data sources had similar predictive capacity (R2 = 0.27). Further, apart from mean maximum temperature (TMAX), the two best predictors of flowering time have not commonly been included in phenological models: the number of frost‐free days (NFFD) and the quantity of precipitation as snow (PAS) in the seasons preceding flowering. By vetting these models across an unprecedented number of taxa, this work demonstrates a new approach to phenological modeling. This study harnesses 894,392 digital herbarium records and 1,959 in situ observations to produce the first assessment of the effects of a 25 of climate parameters on the flowering time of 2,468 of angiosperm taxa throughout North using a regression approach—elastic net regularization—that has not previously been used in phenological modeling, but exhibits several advantages over ordinary least squares and stepwise regression. Multivariate models based on herbarium‐derived data and in situ phenological observations had similar predictive capacity. By vetting these models across an unprecedented number of taxa, this work demonstrates a new approach to phenological modeling. [ABSTRACT FROM AUTHOR]

The timing of the seasonal activity of organisms is a tractable indicator of climate change. Many studies in North America have investigated the role of temperature on the onset date of phenological transitions in temperate deciduous trees and found that the onset of leafing and flowering in numerous species has occurred earlier in recent years, apparently in response to higher temperatures in winter and spring., Few studies have examined the climatic and biogeographic drivers of phenological responses in water-limited ecosystems or explored interspecific variation in responses of phenological metrics other than the timing of onset, such as the periodicity or duration of phenological activity., This study used phenological observations of four species of Quercus contributed to the USA National Phenology Network database from 2009 to 2014 to investigate how responses to climate (temperature and precipitation) and geographic location (latitude, longitude and elevation) varied among two western North American species ( Q. agrifolia and Q. lobata) and two eastern and central North American species ( Q. alba and Q. rubra)., Within years, in species in the western, water-limited ecosystems, the phenological phases observed here (bud break, flowers or flower buds) tend to occur intermittently throughout the growing season, and each event is of longer duration than the same phenophases of the temperate-zone species, rendering a single onset date an incomplete metric with which to track responsiveness or to compare species. By contrast, the eastern/central U.S. species were phenologically more responsive than the western species to spatial and temporal variation in winter, spring, and fall precipitation and maximum temperature., Synthesis. Within and between regions these congeners exhibited a diversity of responses to seasonal temperature and precipitation. This indicates that for predictive model development it is critical to understand how each underlying driver influences species that are adapted to different climatic regimes. These results underscore the value of studying a range of phenological metrics and species from a variety of ecosystems to better predict phenological responses to short-term variation and to long-term change in climate. [ABSTRACT FROM AUTHOR]

In terrestrial ecosystems, atmospheric nitrogen (N) deposition has greatly increased N availability relative to other elements, particularly phosphorus (P). Alterations in the availability of N relative to P can affect plant growth rate and functional traits, as well as resource allocation to above- versus belowground biomass ( MA and MB). Biomass allocation among individual plants is broadly size-dependent, and this can often be described as an allometric relationship between MA and MB, as represented by the equation [ABSTRACT FROM AUTHOR]

In habitats where resource availability declines during the growing season , selection may favor early-flowering individuals. Under such ephemerally favorable conditions, late-blooming species (and individuals) may be particularly vulnerable to resource limitation of seed production. In California, a region prone to seasonal drought, members of the annual genus Clarkia are among the last to flower in the spring. We compared pollen limitation ( PL) of seed set and outcrossing rates between early- and late-flowering individuals in two mixed-mating Clarkia taxa to detect whether flowering time is associated with changes in seed set due to resource depletion, PL, or increased selfing. In 2008-2010, we hand-pollinated one flower on a total of 1855 individual plants either Early (near the onset of flowering) or Late (near the end of flowering) in the flowering season and compared seed set to adjacent, open-pollinated flowers on the same stem. To assess the contribution of pollen quality to reproduction, we first (2008) used allozymes to estimate outcrossing rates of seeds produced by Early and Late open-pollinated flowers. Second (2009), we conducted an anther-removal experiment to estimate self-pollen deposition. Seed set in Clarkia unguiculata was not pollen-limited. Clarkia xantiana ssp . xantiana was pollen-limited in 2008 and 2010, but not 2009. PL did not differ between Early and Late treatments. In both taxa, seed set of Early flowers was greater than Late flowers, but not due to PL in the latter. Reproduction was generally pollinator-dependent. Most pollen deposition was xenogamous, and outcrossing rates were >0.7 - and similar between Early and Late periods. These results suggest that pollen receipt and pollen quality remain seasonally consistent. By contrast, the resources necessary to provision seeds decline, reducing the fitness benefits associated with resource allocation to ovules. [ABSTRACT FROM AUTHOR]

For most species, a precise understanding of how climatic parameters determine the timing of seasonal life cycle stages is constrained by limited long-term data. Further, most long-term studies of plant phenology that have examined relationships between phenological timing and climate have been local in scale or have focused on single climatic parameters. Herbarium specimens, however, can expand the temporal and spatial coverage of phenological datasets., Using Trillium ovatum specimens collected over > 100 yr across its native range, we analyzed how seasonal climatic conditions (mean minimum temperature ( Tmin), mean maximum temperature and total precipitation ( PPT)) affect flowering phenology. We then examined long-term changes in climatic conditions and in the timing of flowering across T. ovatum's range., Warmer Tmin advanced flowering, whereas higher PPT delayed flowering. However, Tmin and PPT were shown to interact: the advancing effect of warmer Tmin was strongest where PPT was highest, and the delaying effect of higher PPT was strongest where Tmin was coldest. The direction of temporal change in climatic parameters and in the timing of flowering was dependent on geographic location. Tmin, for example, decreased across the observation period in coastal regions, but increased in inland areas., Our results highlight the complex effects of climate and geographic location on phenology. [ABSTRACT FROM AUTHOR]

Phenology is an integrative science that comprises the study of recurring biological activities or events. In an era of rapidly changing climate, the relationship between the timing of those events and environmental cues such as temperature, snowmelt, water availability, or day length are of particular interest. This article provides an overview of the observer-based plant phenology sampling conducted by the U.S. National Ecological Observatory Network ( NEON), the resulting data, and the rationale behind the design. Trained technicians will conduct regular in situ observations of plant phenology at all terrestrial NEON sites for the 30-yr life of the observatory. Standardized and coordinated data across the network of sites can be used to quantify the direction and magnitude of the relationships between phenology and environmental forcings, as well as the degree to which these relationships vary among sites, among species, among phenophases, and through time. Vegetation at NEON sites will also be monitored with tower-based cameras, satellite remote sensing, and annual high-resolution airborne remote sensing. Ground-based measurements can be used to calibrate and improve satellite-derived phenometrics. NEON's phenology monitoring design is complementary to existing phenology research efforts and citizen science initiatives throughout the world and will produce interoperable data. By collocating plant phenology observations with a suite of additional meteorological, biophysical, and ecological measurements (e.g., climate, carbon flux, plant productivity, population dynamics of consumers) at 47 terrestrial sites, the NEON design will enable continental-scale inference about the status, trends, causes, and ecological consequences of phenological change. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: One proposed function of long styles is to intensify selection among male gametophytes relative to short styles. If so, given sufficient competition, longer styles will have higher rates of pollen tube attritin (failure to reach the style base) with in the style than shorter ones. Alternatively, style length may influence pollen receipt, which itself may affect attrition rates. METHODS: We tested these predictions by collecting senescing styles from wild populations of two insect-pollinated Clarkia species. We examined the number of pollen grains adhering to the stigma, length of styles, and rates of attrition from the stigma surface to the stigma-style junction (SSJ), from the SSJ to the style base, and from the stigma surface to the style base. Multivariate analyses estimated the independent effects of pollen grains per stigma, the number of pollen tubes at the SSJ, and style length on attrition. KEY RESULTS: Style length was generally positively correlated w ith pollen receipt, and the number of pollen grains per stigma was positively correlated with all three attrition rates. In neither species was any attrition rate affected by style length independent of the number of pollen grains per stigma. CONCLUSIONS: Pollen attrition was mediated by style length, but the function of style length was primarily to increase the number of germinating pollen grains, which affected attrition rates either through stigma clogging or pollen-pollen interactions. Style length may have a direct effect on pollen receipt due to the stigma's position relative to pollinator body parts, but traits correlated with style length may also directly affect pollen receipt. [ABSTRACT FROM AUTHOR]

Ecologists and evolutionary biologists have been interested in the functional biology of pollen since the discovery in the 1800s that pollen grains encompass tiny plants (male gametophytes) that develop and produce sperm cells. After the discovery of double fertilization in flowering plants, botanists in the early 1900s were quick to explore the effects of temperature and maternal nutrients on pollen performance, while evolutionary biologists began studying the nature of haploid selection and pollen competition. A series of technical and theoretic developments have subsequently, but usually separately, expanded our knowledge of the nature of pollen performance and how it evolves. Today, there is a tremendous diversity of interests that touch on pollen performance, ranging from the ecological setting on the stigma, structural and physiological aspects of pollen germination and tube growth, the form of pollen competition and its role in sexual selection in plants, virus transmission, mating system evolution, and inbreeding depression. Given the explosion of technical knowledge of pollen cell biology, computer modeling, and new methods to deal with diversity in a phylogenetic context, we are now more than ever poised for a new era of research that includes complex functional traits that limit or enhance the evolution of these deceptively simple organisms. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: Climate change models for California predict a warmer, drier future, potentially resulting in shorter growing seasons. If phenotypic differences between closely related species currently distributed across a moisture and temperature gradient represent adaptations to their abiotic environment, then as conditions become warmer and drier, populations presently adapted to cooler and wetter conditions may evolve to become more similar to those adapted to warmer and drier conditions. Two sister species, Clarkia unguiculata and C exilis, are distributed across a moisture and temperature gradient in the southern Sierra Nevada, providing an opportunity to predict how this process may occur. METHODS: In a greenhouse experiment using wild-collected seeds from 11 populations in the southern Sierra Nevada, we examined relationships among elevation, climatic conditions, and population means for each trait, then evaluated bivariate relationships among maternal family means, using raw values and controlling for population and seed mass effects on phenotype. KEY RESULTS: Clarkia exilis occupied warmer, drier conditions, typically at lower elevations, than C. unguiculata did and flowered earlier and faster, producing smaller flowers with lower herkogamy. In C. unguiculata, petal area, herkogamy, and the rate of flower production were positively correlated with days tofirst flower. CONCLUSIONS: If selection favors earlier flowering, smaller petals, or faster flower production in C unguiculata, then the genetic correlations among these traits should reinforce their joint evolution. Moreover, the correlations between these traits and herkogamy may promote the evolution of self-fertilization as an indirect response to selection, a previously unrecognized potential outcome of climate change. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: Project Baseline is a seed bank that offers an unprecedented opportunity to examine spatial and temporal dimensions of micro-evolution during an era of rapid environmental change. Over the upcoming 50 years, biologists will withdraw genetically representative samples of past populations from this time capsule of seeds and grow them contemporaneously with modern samples to detect any phenotypic and molecular evolution that has occurred during the intervening time. METHODS: We carefully developed this living genome bank using protocols to enhance its experimental value by collecting from multiple populations and species across a broad geographical range in sites that are likely to be preserved into the future. Seeds are accessioned with site and population data and are stored by maternal line under conditions that maximize seed longevity. This open-access resource will be available to researchers at regular intervals to evaluate contemporary evolution. KEY RESULTS: To date, the Project Baseline collection includes 100-200 maternal lines of each of 61 species collected from over 831 populations on sites that are likely to be preserved into the future across the United States (∼78,000 maternal lines). Our strategically designed collection circumvents some problems that can cloud the results of "resurrection" studies involving naturally preserved or existing seed collections that are available fortuitously. CONCLUSIONS: The resurrection approach can be coupled with long-established and newer techniques over the next five decades to elucidate genetic change and thereby vastly improve our understanding of temporal and spatial changes in phenotype and the evolutionary processes underlying it. [ABSTRACT FROM AUTHOR]

* Premise of the study: One explanation for the evolution of selfing, the drought escape hypothesis, proposes that self-fertilization may evolve under conditions of intensifying seasonal drought as part of a suite of traits that enable plants to accelerate the completion of their life cycle, thereby escaping late-season drought. Here, we test two fundamental assumptions of this hypothesis in Clarkia xantiana: (1) that a seasonal decline in precipitation causes an increase in drought stress and (2) that this results in changes in physiological performance, reflecting these deteriorating conditions. * Methods: We examined seasonal and interannual variation in abiotic environmental conditions (estimated by ambient temperature, relative humidity, predawn leaf water potentials, and carbon isotope ratios) and physiological traits (photosynthesis, conductance, transpiration, instantaneous water-use efficiency, ascorbate peroxidase and glutathione reductase activities, quantum yield of photosystem II, PSII potential efficiency) in field populations of C. xantiana in 2009 and 2010. * Key results: In both years, plants experienced intensifying drought across the growing season. Gas exchange rates decreased over the growing season and were lower in 2009 (a relatively dry year) than in 2010, suggesting that the temporal changes from early to late spring were directly linked to the deteriorating environmental conditions. * Conclusions: Seasonal declines in transpiration rate may have increased survival by protecting plants from desiccation. Concomitant declines in photosynthetic rate likely reduced the availability of resources for seed production late in the season. Thus, the physiological patterns observed are consistent with the conditions required for the drought escape hypothesis. [ABSTRACT FROM AUTHOR]

Phenological events - defined points in the life cycle of a plant or animal - have been regarded as highly plastic traits, reflecting flexible responses to various environmental cues., The ability of a species to track, via shifts in phenological events, the abiotic environment through time might dictate its vulnerability to future climate change. Understanding the predictors and drivers of phenological change is therefore critical., Here, we evaluated evidence for phylogenetic conservatism - the tendency for closely related species to share similar ecological and biological attributes - in phenological traits across flowering plants. We aggregated published and unpublished data on timing of first flower and first leaf, encompassing ˜4000 species at 23 sites across the Northern Hemisphere. We reconstructed the phylogeny for the set of included species, first, using the software program Phylomatic, and second, from DNA data. We then quantified phylogenetic conservatism in plant phenology within and across sites., We show that more closely related species tend to flower and leaf at similar times. By contrasting mean flowering times within and across sites, however, we illustrate that it is not the time of year that is conserved, but rather the phenological responses to a common set of abiotic cues., Our findings suggest that species cannot be treated as statistically independent when modelling phenological responses., Synthesis. Closely related species tend to resemble each other in the timing of their life-history events, a likely product of evolutionarily conserved responses to environmental cues. The search for the underlying drivers of phenology must therefore account for species' shared evolutionary histories. [ABSTRACT FROM AUTHOR]

Premise of the study: Numerous long-term studies in seasonal habitats have tracked interannual variation in first flowering date (FFD) in relation to climate, documenting the effect of warming on the FFD of many species. Despite these efforts, long-term phenological observations are still lacking for many species. If we could forecast responses based on taxonomic affinity, however, then we could leverage existing data to predict the climate-related phenological shifts of many taxa not yet studied. Methods: We examined phenological time series of 1226 species occurrences (1031 unique species in 119 families) across seven sites in North America and England to determine whether family membership (or family mean FFD) predicts the sensitivity of FFD to standardized interannual changes in temperature and precipitation during seasonal periods before flowering and whether families differ significantly in the direction of their phenological shifts. Key results: Patterns observed among species within and across sites are mirrored among family means across sites; early-flowering families advance their FFD in response to warming more than late-flowering families. By contrast, we found no consistent relationships among taxa between mean FFD and sensitivity to precipitation as measured here. Conclusions: Family membership can be used to identify taxa of high and low sensitivity to temperature within the seasonal, temperate zone plant communities analyzed here. The high sensitivity of early-flowering families (and the absence of early-flowering families not sensitive to temperature) may reflect plasticity in flowering time, which may be adaptive in environments where early-season conditions are highly variable among years. [ABSTRACT FROM AUTHOR]

The use of local seed sources for revegetation is accepted practice to reduce the potential that propagules will be poorly adapted to site conditions. However, data are often lacking to determine the distance within which seed sources represent local genotypes. Short-term reciprocal transplant studies represent a class of tools to detect local adaptation of target species. We conducted a reciprocal transplant of Nassella pulchra between two central California locations to test for adaptation to local environmental conditions over a 3-year period. Experimental plots at one location were split between grazed and ungrazed sites to evaluate the potential influence of livestock grazing on the detection or magnitude of local adaptation. During each year of the study, evidence of a home-site advantage depended on the location, traits studied, and population. At the end of the 3-year study period, however, we detected consistent evidence of a home-site advantage for seedling biomass among grazed sites at one location and ungrazed plots at the other location. In effect, local adaptation was only apparent in the final year of the study. Short-term reciprocal transplant studies are an effective tool to guide the selection of seed sources most likely to germinate and to become established at a restoration site, but such studies cannot rule out local adaptation, which may not be immediately detectable. [ABSTRACT FROM AUTHOR]

Ecology Letters (2012) Abstract Shifts in species' phenology in response to climate change have wide-ranging consequences for ecological systems. However, significant variability in species' responses, together with limited data, frustrates efforts to forecast the consequences of ongoing phenological changes. Herein, we use a case study of three North American plant communities to explore the implications of variability across levels of organisation (within and among species, and among communities) for forecasting responses to climate change. We show how despite significant variation among species in sensitivities to climate, comparable patterns emerge at the community level once regional climate drivers are accounted for. However, communities differ with respect to projected patterns of divergence and overlap among their species' phenological distributions in response to climate change. These analyses and a review of hypotheses suggest how explicit consideration of spatial scale and levels of biological organisation may help to understand and forecast phenological responses to climate change. [ABSTRACT FROM AUTHOR]

* Premise of the study: The evolution of self-fertilization often occurs in association with other floral, life history, and fitness-related traits. A previous study found that field populations of Clarkia exilis (a predominantly autogamous selfer) and its sister species, Clarkia unguiculata (a facultative outcrosser) differ in mean photosynthetic rates and instantaneous water use efficiency (WUE i). Here, we investigate the strength and direction of selection on these traits in multiple populations of each taxon to determine whether natural selection may contribute to the phenotypic differences between them. * Methods: In spring 2008, we measured instantaneous gas exchange rates in nine populations during vegetative growth (Early) and/or during flowering (Late). We conducted selection gradient analyses and estimated selection differentials within populations and across pooled conspecific populations to evaluate the strength, direction, and consistency of selection on each trait early and late in the season. * Key results: The direction and relative strength of selection on photosynthetic rates in these taxa corresponds to the phenotypic difference between them; C. exilis has higher photosynthetic rates than C. unguiculata, as well as stronger, more consistent selection favoring rapid photosynthesis throughout the growing season. Patterns of selection on transpiration, WUE i , and the timing of flowering progression are less consistent with phenotypic differences (or lack thereof) between taxa. * Conclusions: We detected several examples where selection was consistent with the phenotypic divergence between sister taxa, but there were also numerous instances that were equivocal or in which selection did not predict the realized phenotypic difference between taxa. [ABSTRACT FROM AUTHOR]

Summary 1. Plants produce biomass and then allocate some of it to reproductive structures, so the relationship between reproductive (R) and vegetative (V) mass is a fundamental aspect of a plant's reproductive strategy. 2. Differences among populations or taxa in the allometric relationship between R and V have been attributed to environmental conditions. We hypothesize that populations and taxa living at high elevations should exhibit a lower log R vs. log V slope than those in more benign environments, because the environmental limits on size in alpine environments should favour a relatively large reproductive allocation at smaller sizes and a smaller investment in reproduction per additional unit of biomass accumulated. 3. We investigated variation in the allometric relationship between R and V among 44 naturally occurring populations representing 24 species of Pedicularis in the Tibetan Plateau, to test the hypothesis that the slope of the relationship declines with increasing elevation. 4. There was a significant negative relationship between the slope of the log R vs. log V relationship and elevation among populations, although the relationship among populations within species varied. We interpret this in terms of abiotic limitations on size and decreasing efficiency of resource allocation to reproduction with increasing plant size (measured as vegetative biomass) at high elevations. We also found a significant positive relationship between the y-intercept of the regression of log R on log V and elevation across species, but the relationship was not significant among populations within species. The combination of the lower slopes and the higher y-intercepts for high-elevation populations means that plants growing at high elevations allocate proportionately more biomass to reproduction at smaller sizes and less at larger sizes than plants growing at lower elevations. 5. Synthesis. The allometric slope (exponent) of the R-V relationship decreases with increasing elevation among Pedicularis populations and species, reflecting fundamental changes in the costs and benefits of increased vegetative biomass with elevation. [ABSTRACT FROM AUTHOR]

1. We examined geographic variation in mean individual seed mass (MISM) among 38 populations within and across nine Pedicularis species in the eastern Tibetan Plateau, measuring the effects of one extrinsic factor (elevation) and two intrinsic factors (plant size and mean seed number per fruit). 2. Across all populations, elevation is negatively correlated with MISM; within eight of nine species, this pattern is also observed among populations. This relationship, however, is not significant when controlling for variation in plant size or seeds per fruit. High-elevation populations produce smaller plants with more seeds per fruit than low-elevation populations; controlling for these variables eliminates the negative correlation between elevation and MISM. 3. Across all populations, the predicted trade-off between MISM and seeds per fruit is consistently strong, with or without controlling for the effects of plant size. By contrast, the trade-off between MISM and total seasonal fecundity can be detected only when controlling for plant size. 4. Independent of plant size, populations that produce small seeds do not support individuals with particularly low reproductive yield (fecundity × MISM). Accordingly, high-elevation populations exhibit neither lower reproductive yield nor smaller seeds than expected given their lower biomass. 5. Synthesis. In Pedicularis, elevation, plant size and seeds per fruit are all correlated with MISM among populations across species. Elevation is less important, however, than intrinsic factors in determining the MISM of a population; the effect of elevation on MISM disappears when the effects of intrinsic factors are controlled statistically. The observed decline in MISM with increasing elevation is therefore partly mediated by the decline in plant size and partly by an increase in mean seed number per fruit with elevation. Altitudinal variation in MISM across populations or species has been described before, but this is the first study to control for the effect of intrinsic factors simultaneously. This result calls into question the conclusions of studies that have detected geographic variation in MISM without controlling for variation in intrinsic factors. [ABSTRACT FROM AUTHOR]

• Premise of the study: The study of geographic variation in ecologically important traits within and among taxa is a first step toward understanding the environmental factors that contribute to population differentiation and species divergence. This study examines variation in mean sex allocation per flower (androecium mass/gynoecium mass) among 49 wild populations representing 12 Pedicularis species across an elevation gradient on the eastern Tibetan Plateau. • Methods: We used population means to evaluate sources of variation in per-flower sex allocation within and across species. In particular, we evaluate the relative influence of intrinsic (i.e., plant size, estimated as aboveground stem biomass) vs. extrinsic factors affecting mean sex allocation among populations. • Key results: Mean sex allocation per flower (the relative investment in male floral organs) is negatively correlated with mean plant size; populations of large plants produce relatively female-biased flowers. This relationship between mean plant size and mean sex allocation is not statistically significant, however, when the effect of elevation is controlled statistically. Among populations within and across species, mean sex allocation increases with elevation. This relationship persists even when the effect of mean plant size is controlled statistically. Factors associated with increasing elevation appear to favor genotypes and/ or taxa with male-biased flowers. • Conclusion: Extrinsic environmental conditions may be more important than intrinsic resource status in determining patterns of geographic variation in mean sex allocation among populations or species of Pedicularis. We cannot conclude whether the effect of elevation on mean sex allocation is the result of environmentally induced plasticity, genetically based adaptation, or species sorting, but it is only partly mediated by mean plant size. [ABSTRACT FROM AUTHOR]

The study of plant responses to touch, particularly the responses of leaves, stems, and roots, has a long history. By contrast, floral responses are relatively unexplored. Stigma closure is common in the Bignoniaceae, but the factors influencing it are not well understood. We investigated factors influencing stigma closure and reopening and its effects on pollen receipt in seven Oroxylum indicurn trees near Hat Yai, Thailand. The effects of pressure, conspecific and heterospecific pollen, and pollen load (the amount of pollen deposited) on stigma behavior were examined in 270 flowers (of the total 430 flowers evaluated in the entire study). Pressure alone resulted in faster closure than did conspecific pollination and faster reopening than did heterospecific pollination. Stigmas never reopened after conspecific pollination. Pollen load had no effect on stigma behavior. Stigmas discriminated between conspecific and foreign pollen; they reopened only after pollination with the latter. A manipulative experiment revealed that stigma closure did not affect the number of conspecific pollen grains received. We also counted pollen tubes in styles that were either hand-supplemented with outcross conspecific pollen or open-pollinated. Pollen tube numbers were highest after light pollination (∼900 grains), indicating that interference among pollen grains may occur after pollination with very heavy loads (>6000 grains). Possible fitness consequences of these responses are discussed. [ABSTRACT FROM AUTHOR]

• It has been proposed that natural selection should favor distinct temporal patterns of sex allocation in selfing vs pollinator-dependent taxa. In autogamous selfers in which pollen receipt is highly reliable, selection should favor genotypes that maintain low and stable pollen to ovule (P : O) ratios throughout flowering. By contrast, in outcrossers the optimum P : O ratio of an individual's flowers will depend on pollinator abundances and mating opportunities, both of which may vary over time. In this case, selection may favor temporal variation among flowers in the P : O ratio. An opposing prediction is that selfing taxa will be developmentally more unstable than outcrossers because of lower homeostasis caused by high homozygosity. • We compared temporal changes in the P : O ratio in two pairs of sister taxa in the genus Clarkia. We examined hundreds of glasshouse-raised maternal families representing three wild populations each of the outcrossing, insect-pollinated Clarkia unguiculata, the facultatively autogamous Clarkia exilis and the outcrossing and selfing subspecies of Clarkia xantiana: ssp. xantiana and parviflora, respectively. • Temporal change in the P : O ratio was significantly greater in both outcrossers than in their selfing sister taxa, although the proportional changes in the P : O ratio (relative to the first bud produced) did not differ significantly between sister taxa (0.07 <  P < 0.10). • Our results provide partial support for the hypothesis that the P : O ratio is more stable in selfing than in outcrossing taxa and reject the hypothesis that selfers are less stable. [ABSTRACT FROM AUTHOR]

The number of ovules per flower varies over several orders of magnitude among angiosperms. Here we consider evidence that stochastic uncertainty in pollen receipt and ovule fertilization has been a selective factor in the evolution of ovule number per flower. We hypothesize that stochastic variation in floral mating success creates an advantage to producing many ovules per flower because a plant will often gain more fitness from occasional abundant seed production in randomly successful flowers than it loses in resource commitment to less successful flowers. Greater statistical dispersion in pollination and fertilization among flowers increases the frequency of windfall success, which should increase the strength of selection for greater ovule number per flower. We therefore looked for evidence of a positive relationship between ovule number per flower and the statistical dispersion of pollen receipt or seed number per flower in a comparative analysis involving 187 angiosperm species. We found strong evidence of such a relationship. Our results support the hypothesis that unpredictable variation in mating success at the floral level has been a factor in the evolution of ovule packaging in angiosperms. [ABSTRACT FROM AUTHOR]

1. Dioecious species may suffer a reproductive handicap compared to sympatric cosexual (hermaphroditic or monoecious) species of equal abundance simply because populations of dioecious species contain fewer seed-producing individuals. To compensate for this population-level disadvantage, natural selection in dioecious populations should favour females that reallocate those reproductive resources, that in cosexual individuals are invested in male function, towards the production of higher quality or more seeds, potentially leading to a higher density of established individuals. 2. We tested for two compensatory fitness advantages (heavier seeds and higher population densities) in dioecious species in the flora of a high-diversity tropical forest in Western Amazonia. Using a phylogenetically corrected generalized linear modelling approach, we examined the phylogenetic dependence (Pagel's λ) of seed mass, abundance, growth form and wood-specific gravity for 641 species from Yasuní National Park, Ecuador. Seed mass was highly correlated with phylogeny, but abundance was not. We then used AIC values to test for models (including breeding system) that best explained seed mass and species abundance, using the maximum likelihood values of λ. 3. Contrary to expectations, dioecious species exhibited no advantage with respect to either seed mass or population density; there was no difference in mean seed mass or abundance between dioecious species and their most closely related cosexual species. 4. We compared our results with those from a similar forest in the Tambopata Wildlife Reserve, Peru. There, dioecious shrubs had larger seeds than hermaphroditic shrubs, but (as for Yasuní) there was no difference between breeding systems in seed size among trees. Differences in the composition of species sampled may have contributed to differences in the results between the two sites. 5. We suggest that other individual fitness advantages, such as reduced inbreeding, increased dispersal, superior seed quality (e.g. better defences) or increased seed number contribute to the persistence of dioecious populations at Yasuní. 6. Synthesis. Similarities in the statistical models of seed mass and abundance for plant species in a Western Amazonian rain forest indicate that selective pressures on the seed-producing individuals are similar, despite variation in sexual system. Therefore, breeding system may play a more limited role in community assembly of tropical forests than was previously considered. [ABSTRACT FROM AUTHOR]

Multiple field populations of two pairs of diploid sister taxa with contrasting mating systems in the genus Clarkia (Onagraceae) were surveyed to test predictions concerning the effects of resource status, estimated as plant size, on pollen and ovule production and on the pollen:ovule (P:O) ratio of flowers. Most theoretical models of size-dependent sex allocation predict that, in outcrossing populations, larger plants should allocate more resources to female function. Lower P:O ratios in larger plants compared to smaller plants have been interpreted as supporting this prediction. In contrast, we predicted that P:O ratio should not vary with plant size in predominantly selfing plants, in which each flower contributes to reproductive success equally through male and female function. We found that, in all four taxa, both ovule and pollen production per flower usually increased significantly with plant size and that the shape of this relationship was decelerating. However, ovule production either decelerated more rapidly than or at the same rate as pollen production with plant size. Consequently,the P:O ratio increased or had no relationship with plant size. This relationship was population-specific (not taxon-specific) and independent of the mating system. Possible explanations for the increasing maleness with plant size are discussed. [ABSTRACT FROM AUTHOR]

A well-known, but largely untested, prediction in plant reproductive ecology is that dioecious taxa should produce larger, more, higher-quality, or better-defended seeds than cosexual taxa. Using a data set composed of 972 species in 104 families, representing the flora of the Tambopata Wildlife Reserve (Madre de Dios, Peru), we evaluated the first component of this prediction, examining ecological and evolutionary relationships between breeding system and mean seed size with two kinds of tests. First, we conducted cross-species analyses to determine whether species with different breeding systems differed significantly with respect to mean individual seed size. Second, we used a hypothesized phylogeny to identify pairs of the most closely related taxa or clades within the Tambopata community that differed with respect to breeding system. Comparing pair members allowed us to determine whether evolutionary divergence in breeding system (between taxa with unisexual vs. cosexual individuals) was consistently associated with evolutionary change in seed size. In both analyses, we controlled for potentially confounding effects of growth form by examining these relationships within woody and nonwoody taxa. Cross-species analyses revealed that dioecious species produced larger seeds than cosexual species among woody species, shrubs, lianas (each growth form analyzed separately), and all species pooled, but not among trees. Phylogenetically independent contrasts upheld the significant association between breeding system and seed size among woody taxa, lianas, and all taxa pooled, but not among shrubs. We discuss the implications of our findings for evolutionary hypotheses regarding associations between dioecy and seed size. [ABSTRACT FROM AUTHOR]

• Plants growing at low density can suffer from Allee effects as a result of pollen limitation. Previous studies of Allee effects have focused on the effects of variation among populations in size or density on reproduction. Here, the effects of plant distribution within populations on fitness components are explored in a rare plant, Aconitum napellus ssp. lusitanicum, and ecological and genetic mechanisms underlying these effects are identified. • To detect pollen limitation, seed production was compared under natural versus hand-supplemented pollinations on inflorescences of different sizes in natural patches differing both in flower density and in isolation from other patches. Germination rate and juvenile survival of seeds produced in low- and high-density patches were also compared. • Pollen-supplemented flowers always produced more seeds than open-pollinated flowers, especially among small plants and plants growing at low density. Offspring produced in low-density patches exhibited lower fitness that those produced in high-density patches. This could have been caused by post-fertilization mechanisms, including inbreeding depression or differential maternal resource allocation. • These results show that Allee effects on fitness components (ecological and genetic Allee effects) occur within A. napellus populations at different spatial scales. The spatial distribution of plants seems to be a crucial factor affecting reproductive output and fitness. [ABSTRACT FROM AUTHOR]

Adaptation to environmental factors may influence the germination and establishment of focal species in ecological restoration. Reciprocal transplants remain one of the best methods to detect local adaptation, but long-term studies are often not feasible. We conducted reciprocal transplants of the native California bunchgrasses Elymus glaucus and Bromus carinatus between two central California locations to seek evidence of adaptation to local environmental conditions in a single growing season. Experimental plots at one location included grazed and ungrazed sites. The combination of locations and grazing treatments allowed us to determine whether the ability to detect evidence for adaptation depended on grazing regime. In addition, we measured the direct effects of grazing on seedling growth and survival concurrent with our investigation of local adaptation. We detected a homesite advantage for seedling growth or survival in both species, but the factors contributing to adaptive differentiation were species specific. Evidence of local adaptation was detected for seedling biomass in Bromus and for survivorship in Elymus. The homesite advantage observed in both species was greatly reduced under grazed conditions and in Elymus was significant only in the ungrazed plots. Climate and soil analyses detected significant differences between locations in five soil attributes and two climate variables. In particular, differences in exchangeable magnesium indicated that one of the two transplant locations consisted of serpentine soil, which is widely known to drive adaptation in plant populations. Together, these results suggest that it is possible to investigate the scale and factors involved in local adaptation with short-term transplant studies. [ABSTRACT FROM AUTHOR]

Autogamously self-fertilizing taxa have evolved from outcrossing progenitors at least 12 times in the annual wildflower genus, Qarkia (Onagraceae). In C. xantiana, individuals of the selfing subspecies (ssp. parviflora) flower at an earlier age, produce successive flowers more rapidly, and produce flowers that complete their development more rapidly than their outcrossing counterparts (ssp. xantiana). Two hypotheses have been proposed to explain the joint evolution of these whole-plant and individual floral traits. The accelerated life cycle hypothesis proposes that selection favoring a short life cycle in environments with short growing seasons (such as those typically occupied by parviflora) has independently favored genotypes with early reproduction, synchronous flower production, and rapidly developing, self-fertilizing flowers. The correlated response to selection hypothesis similarly proposes that selection in environments with short growing seasons favors early reproduction, but that rapid floral development and increased selfing evolve as correlated responses to selection due to genetic linkage (or pleiotropy) affecting both whole-plant and floral development. We conducted a greenhouse experiment using maternal families from two field populations of each subspecies to examine covariation between floral and whole-plant traits within and among populations to seek support for either of these hypotheses. Our results are consistent with the accelerated life cycle hypothesis but not with the correlated response to selection hypothesis. [ABSTRACT FROM AUTHOR]

Determining whether seed production is pollen limited has been an area of intensive empirical study over the last two decades. Yet current evidence does not allow satisfactory assessment of the causes or consequences of pollen limitation. Here, we critically evaluate existing theory and issues concerning pollen limitation. Our main conclusion is that a change in approach is needed to determine whether pollen limitation reflects random fluctuations around a pollen-resource equilibrium, an adaptation to stochastic pollination environments, or a chronic syndrome caused by an environmental perturbation. We formalize and extend D. Haig and M. Westoby's conceptual model, and illustrate its use in guiding research on the evolutionary consequences of pollen limitation, i.e., whether plants evolve or have evolved to ameliorate pollen limitation. This synthesis also reveals that we are only beginning to understand when and how pollen limitation at the plant level translates into effects on plant population dynamics. We highlight the need for both theoretical and empirical approaches to gain a deeper understanding of the importance of life-history characters, Allee effects, and environmental perturbations in population declines mediated by pollen limitation. Lastly, our synthesis identifies a critical need for research on potential effects of pollen limitation at the community and ecosystem levels. [ABSTRACT FROM AUTHOR]

Summary • We studied how the genetic structure helps in understanding the demographic history of a species and predicting its future, using the endangered orchid Spiranthes spiralis . • A Parisian population was mapped during three consecutive years. Its genetic structure was assessed using isozymes and compared with several populations from southern France. Seed dispersal distances were experimentally evaluated. Computer simulations were performed to reconstruct the population history. • Unlike southern France populations, the Parisian population exhibited two genetically distinct patches. Seed dispersal was limited to the neighbourhood of the fruiting plants. The simulations showed that the best explanation for the observed genetic structure was a recent colonization event by two small groups of founders. • Current patterns of gene flow should result in a rapid genetic and demographic homogenization. New populations of S. spiralis could be successfully established by few founders and exhibit significant growth rates without any detrimental effects of inbreeding. This offers insight into the successful management of endangered plant species in urban areas. [ABSTRACT FROM AUTHOR]

Provides information on a study which examined ontogenetic variation in sex allocation among flowers and correlations between size and sex allocation among whole plants of the annual herb Clarkia unguiculata. Methods; Results; Discussion.

Presents information on a study which determined the intensity of cryptic self-incompatibility (CSI) in Clarkia unguiculata from a large population of C. unguiculata in Southern California. Evolution of CSI; Materials and methods of the study; Results and discussion.