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Summary 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. This primer summarizes research on climate change responses for eight intensively studied herbaceous plant species. We establish generalities across species, examine limitations, and propose a path forward. Climate change has reduced fitness, caused maladaptation, and/or led to population declines in at least part of the range of all six forb species. 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 generally widespread, span large elevation ranges, and have substantial genetic and phenotypic variation. The focal species have diverse life histories, reproductive strategies, and habitats, and most are native to North America. Thus, species that are rare, habitat specialists, or endemic to other parts of the world are poorly represented in this review. 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 biotic interactions in climate change responses. 

ABSTRACT Theory suggests that the drivers of demographic variation and local adaptation are shared and may feedback on one other. Despite some evidence for these links in controlled settings, the relationship between local adaptation and demography remains largely unexplored in natural conditions. Using 10 years of demographic data and two reciprocal transplant experiments, we tested predictions about the relationship between the magnitude of local adaptation and demographic variation (population growth rates and their elasticities to vital rates) across 10 populations of a well‐studied annual plant. In both years, we found a strong unimodal relationship between mean home‐away local adaptation and stochastic population growth rates. Other predicted links were either weakly or not supported by our data. Our results suggest that declining and rapidly growing populations exhibit reduced local adaptation, potentially due to maladaptation and relaxed selection, respectively. 

Abstract Plant species’ distributions are often thought to overwhelmingly reflect their climatic niches. However, climate represents only a fraction of then‐dimensional environment to which plant populations adapt, and studies are increasingly uncovering strong effects of nonclimatic factors on species’ distributions. We used a manipulative, factorial field experiment to quantify the effects of soil environment and precipitation (the putatively overriding climatic factor) on plant lifetime fitness outside the geographic range boundary of a native California annual plant. We grew plants outside the range edge in large mesocosms filled with soil from either within or outside the range, and plants were subjected to either a low (ambient) or high (supplemental) spring precipitation treatment. Soil environment had large effects on plant lifetime fitness that were similar in magnitude to the effects of precipitation. Moreover, mean fitness of plants grown with within‐range soil in the low precipitation treatment approximated that of plants grown with beyond‐range soil in the high precipitation treatment. The positive effects of within‐range soil persisted in the second, wetter year of the experiment, though the magnitude of the soil effect was smaller than in the first, drier year. These results are the first we know of to quantify the effects of edaphic variation on plant lifetime fitness outside a geographic range limit and highlight the need to include factors other than climate in models of species’ distributions. 

Summary Interactions between plants and soil fungi and bacteria are ubiquitous and have large effects on individual plant fitness. However, the degree to which spatial variation in soil microbial communities modulates plant species’ distributions remains largely untested.Using the California native plantClarkia xantianassp.xantianawe paired glasshouse and field reciprocal transplants of plant populations and soils to test whether plant–microbe interactions affect the plant’s geographic range limit and whether there is local adaptation between plants and soil microbe communities.In the field and glasshouse, one of the two range interior inocula had a positive effect on plant fitness. In the field, this benefit was especially pronounced at the range edge and beyond, suggesting possible mutualist limitation. In the glasshouse, soil inocula from beyond‐range tended to increase plant growth, suggesting microbial enemy release beyond the range margin. Amplicon sequencing revealed stark variation in microbial communities across the range boundary.Plants dispersing beyond their range limit are likely to encounter novel microbial communities. InC. x. xantiana, our results suggest that range expansion may be facilitated by fewer pathogens, but could also be hindered by a lack of mutualists. Both negative and positive plant–microbe interactions will likely affect contemporary range shifts. 

Introgression is pervasive across the tree of life but varies across taxa, geography, and genomic regions. However, the factors modulating this variation and how they may be affected by global change are not well understood. Here, we used 200 genomes and a 15-y site-specific environmental dataset to investigate the effects of environmental variation and mating system divergence on the magnitude of introgression between a recently diverged outcrosser-selfer pair of annual plants in the genusClarkia. These sister taxa diverged very recently and subsequently came into secondary sympatry where they form replicated contact zones. Consistent with observations of other outcrosser-selfer pairs, we found that introgression was asymmetric between taxa, with substantially more introgression from the selfer to the outcrosser. This asymmetry was caused by a bias in the direction of initial F1 hybrid formation and subsequent backcrossing. We also found extensive variation in the outcrosser’s admixture proportion among contact zones, which was predicted nearly entirely by interannual variance in spring precipitation. Greater fluctuations in spring precipitation resulted in higher admixture proportions, likely mediated by the effects of spring precipitation on the expression of traits that determine premating reproductive isolation. Climate-driven hybridization dynamics may be particularly affected by global change, potentially reshaping species boundaries and adaptation to novel environments. 

Bet hedging consists of life history strategies that buffer against environmental variability by trading off immediate and long-term fitness. Delayed germination in annual plants is a classic example of bet hedging and is often invoked to explain low germination fractions. We examined whether bet hedging explains low and variable germination fractions among 20 populations of the winter annual plant Clarkia xantiana ssp. xantiana that experience substantial variation in reproductive success among years. Leveraging 15 years of demographic monitoring and 3 years of field germination experiments, we assessed the fitness consequences of seed banks and compared optimal germination fractions from a density-independent bet-hedging model to observed germination fractions. We did not find consistent evidence of bet hedging or the expected trade-off between arithmetic and geometric mean fitness, although delayed germination increased long-term fitness in 7 of 20 populations. Optimal germination fractions were two to five times higher than observed germination fractions, and among-population variation in germination fractions was not correlated with risks across the life cycle. Our comprehensive test suggests that bet hedging is not sufficient to explain the observed germination patterns. Understanding variation in germination strategies will likely require integrating bet hedging with complementary forces shaping the evolution of delayed germination. 

Abstract Premise Pollen movement is a crucial component of dispersal in seed plants. Although pollen dispersal is well studied, methodological constraints have made it challenging to directly track pollen flow within multiple populations across landscapes. We labeled pollen with quantum dots, a new technique that overcomes past limitations, to evaluate the spatial scale of pollen dispersal and its relationship with conspecific density within 11 populations of Clarkia xantiana subsp. xantiana , a bee‐pollinated annual plant. Methods We used experimental arrays in two years to track pollen movement across distances of 5–35 m within nine populations and across distances of 10–70 m within two additional populations. We tested for distance decay of pollen dispersal, whether conspecific density modulated dispersal distance, and whether dispersal kernels varied among populations across an environmentally complex landscape. Results Labeled pollen receipt did not decline with distance over 35 m within eight of nine populations or over 70 m within either of two populations. Pollen receipt increased with conspecific density. Overall, dispersal kernels were consistent across populations. Conclusions The surprising uniformity in dispersal distance within different populations was likely influenced by low precipitation and plant density in our study years. This suggests that spatiotemporal variation in the abiotic environment substantially influences the extent of gene flow within and among populations. 

Populations often vary in their evolutionary responses to a shared environmental perturbation. A key hurdle in building more predictive models of rapid evolution is understanding this variation—why do some populations and traits evolve while others do not? We combined long-term demographic and environmental data, estimates of quantitative genetic variance components, a resurrection experiment and individual-based evolutionary simulations to gain mechanistic insights into contrasting evolutionary responses to a severe multi-year drought. We examined five traits in two populations of a native California plant, Clarkia xantiana , at three time points over 7 years. Earlier flowering phenology evolved in only one of the two populations, though both populations experienced similar drought severity and demographic declines and were estimated to have considerable additive genetic variance for flowering phenology. Pairing demographic and experimental data with evolutionary simulations suggested that while seed banks in both populations likely constrained evolutionary responses, a stronger seed bank in the non-evolving population resulted in evolutionary stasis. Gene flow through time via germ banks may be an important, underappreciated control on rapid evolution in response to extreme environmental perturbations.