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Herbaceous plant species have been the focus of extensive, long-term research into climate change responses, but there has been little effort to synthesize results and predicted outlooks from different model species. We summarize research on climate change responses for eight intensively-studied herbaceous plant species. We establish generalities across species, examine limitations, interrogate biases, and propose a path forward. All six forb species exhibit reduced fitness, maladaptation, and/or population declines in at least part of the range. Plasticity alone is likely not sufficient to allow adjustment to shifting climates. Most model species also have spatially-restricted dispersal that may limit genetic and evolutionary rescue. These results are surprising, given that these species are widespread, span large elevation ranges, and generally have substantial levels of genetic and phenotypic variation. The focal species have diverse life histories, reproductive strategies, and habitats, but most are native to North America. Thus, these species may poorly represent rare species, habitat specialists, or species endemic to other parts of the world. We encourage researchers to design demographic and field experiments that evaluate plant traits and fitness in contemporary and potential future conditions across the full life cycle, and that consider the effects of climate change on biotic interactions. 

Abstract In ecological speciation, incipient species diverge due to natural selection that is ecologically based. In flowering plants, different pollinators could mediate that selection (pollinator-mediated divergent selection) or other features of the environment that differ between habitats of 2 species could do so (environment-mediated divergent selection). Although these mechanisms are well understood, they have received little rigorous testing, as few studies of divergent selection across sites of closely related species include both floral traits that influence pollination and vegetative traits that influence survival. This study employed common gardens in sites of the 2 parental species and a hybrid site, each containing advanced generation hybrids along with the parental species, to test these forms of ecological speciation in plants of the genus Ipomopsis. A total of 3 vegetative traits (specific leaf area, leaf trichomes, and photosynthetic water-use efficiency) and 5 floral traits (corolla length and width, anther insertion, petal color, and nectar production) were analyzed for impacts on fitness components (survival to flowering and seeds per flower, respectively). These traits exhibited strong clines across the elevational gradient in the hybrid zone, with narrower clines in theory reflecting stronger selection or higher genetic variance. Plants with long corollas and inserted anthers had higher seeds per flower at the Ipomopsis tenuituba site, whereas selection favored the reverse condition at the Ipomopsis aggregata site, a signature of divergent selection. In contrast, no divergent selection due to variation in survival was detected on any vegetative trait. Selection within the hybrid zone most closely resembled selection within the I. aggregata site. Across traits, the strength of divergent selection was not significantly correlated with width of the cline, which was better predicted by evolvability (standardized genetic variance). These results support the role of pollinator-­mediated divergent selection in ecological speciation and illustrate the importance of genetic variance in determining divergence across hybrid zones. 

Research on floral volatiles has grown substantially in the last 20 years, which has generated insights into their diversity and prevalence. These studies have paved the way for new research that explores the evolutionary origins and ecological consequences of different types of variation in floral scent, including community-level, functional, and environmentally induced variation. However, to address these types of questions, novel approaches are needed that can handle large sample sizes, provide quality control measures, and make volatile research more transparent and accessible, particularly for scientists without prior experience in this field. Drawing upon a literature review and our own experiences, we present a set of best practices for next-generation research in floral scent. We outline methods for data collection (experimental designs, methods for conducting field collections, analytical chemistry, compound identification) and data analysis (statistical analysis, database integration) that will facilitate the generation and interpretation of quality data. For the intermediate step of data processing, we created the R package bouquet , which provides a data analysis pipeline. The package contains functions that enable users to convert chromatographic peak integrations to a filtered data table that can be used in subsequent statistical analyses. This package includes default settings for filtering out non-floral compounds, including background contamination, based on our best-practice guidelines, but functions and workflows can be easily customized as necessary. Next-generation research into the ecology and evolution of floral scent has the potential to generate broadly relevant insights into how complex traits evolve, their genomic architecture, and their consequences for ecological interactions. In order to fulfill this potential, the methodology of floral scent studies needs to become more transparent and reproducible. By outlining best practices throughout the lifecycle of a project, from experimental design to statistical analysis, and providing an R package that standardizes the data processing pipeline, we provide a resource for new and seasoned researchers in this field and in adjacent fields, where high-throughput and multi-dimensional datasets are common. 

Climate change is likely to alter both flowering phenology and water availability for plants. Either of these changes alone can affect pollinator visitation and plant reproductive success. The relative impacts of phenology and water, and whether they interact in their impacts on plant reproductive success remain, however, largely unexplored. We manipulated flowering phenology and soil moisture in a factorial experiment with the subalpine perennial Mertensia ciliata (Boraginaceae). We examined responses of floral traits, floral abundance, pollinator visitation, and composition of visits by bumblebees vs. other pollinators. To determine the net effects on plant reproductive success, we also measured seed production and seed mass. Reduced water led to shorter, narrower flowers that produced less nectar. Late flowering plants produced fewer and shorter flowers. Both flowering phenology and water availability influenced pollination and reproductive success. Differences in flowering phenology had greater effects on pollinator visitation than did changes in water availability, but the reverse was true for seed production and mass, which were enhanced by greater water availability. The probability of receiving a flower visit declined over the season, coinciding with a decline in floral abundance in the arrays. Among plants receiving visits, both the visitation rate and percent of non-bumblebee visitors declined after the first week and remained low until the final week. We detected interactions of phenology and water on pollinator visitor composition, in which plants subject to drought were the only group to experience a late-season resurgence in visits by solitary bees and flies. Despite that interaction, net reproductive success measured as seed production responded additively to the two manipulations of water and phenology. Commonly observed declines in flower size and reward due to drought or shifts in phenology may not necessarily result in reduced plant reproductive success, which in M. ciliata responded more directly to water availability. The results highlight the need to go beyond studying single responses to climate changes, such as either phenology of a single species or how it experiences an abiotic factor, in order to understand how climate change may affect plant reproductive success. 

How climate change influences the dynamics of plant populations is not well understood, as few plant studies have measured responses of vital rates to climatic variables and modeled the impact on population growth. The present study used 25 y of demographic data to analyze how survival, growth, and fecundity respond to date of spring snowmelt for a subalpine plant. Fecundity was estimated by seed production (over 15 y) and also divided into flower number, fruit set, seeds per fruit, and escape from seed predation. Despite no apparent effects on flower number, plants produced more seeds in years with later snowmelt. Survival and probability of flowering were reduced by early snowmelt in the previous year. Based on demographic models, earlier snowmelt with warming is expected to lead to negative population growth, driven especially by changes in seedling establishment and seed production. These results provide a rare example of how climate change is expected to influence the dynamics of a plant population. They furthermore illustrate the potential for strong population impacts even in the absence of more commonly reported visual signs, such as earlier blooming or reduced floral display in early melting years.