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Abstract PremiseSeed germination involves risk; post‐germination conditions might not allow survival and reproduction. Variable, stressful environments favor seeds with germination that avoids risk (e.g., germination in conditions predicting success), spreads risk (e.g., dormancy), or escapes risk (e.g., rapid germination). Germination studies often investigate trait correlations with climate features linked to variation in post‐germination reproductive success. Rarely are long‐term records of population reproductive success available. MethodsSupported by demographic and climate monitoring, we analyzed germination in the California winter‐annualClarkia xantianasubsp.xantiana. Sowing seeds of 10 populations across controlled levels of water potential and temperature, we estimated temperature‐specific base water potential for 20% germination, germination time weighted by water potential above base (hydrotime), and a dormancy index (frequency of viable, ungerminated seeds). Mixed‐effects models analyzed responses to (1) temperature, (2) discrete variation in reproductive success (presence or absence of years with zero seed production by a population), and (3) climate covariates, mean winter precipitation and coefficient of variation (CV) of spring precipitation. For six populations, records enabled analysis with a continuous metric of variable reproduction, the CV of per‐capita reproductive success. ResultsPopulations with more variable reproductive success had higher base water potential and dormancy. Higher base water potential and faster germination occurred at warmer experimental temperatures and in seeds of populations with wetter winters. ConclusionsGeographic variation in seed germination in this species suggests local adaptation to demographic risk and rainfall. High base water potential and dormancy may concentrate germination in years likely to allow reproduction, while spreading risk among years. 

Abstract Background and AimsPollination failure occurs from insufficient pollen quantity or quality. However, the relative contributions of pollen quantity vs. quality to overall pollen limitation, and how this is affected by the co-flowering context, remain unknown for most plant populations. Here, we studied patterns of pollen deposition and pollen tube formation across populations of four predominately outcrossing species in the genus Clarkia to evaluate how the richness of co-flowering congeners affects the contribution of pollen quantity and quality to pollen limitation. MethodsWe partition variation in pollen deposition and pollen tube production across individuals, populations and species to identify the main sources of variation in components of reproductive success. We further quantify the relative contribution of pollen quantity and quality limitation to the reproductive success of the four Clarkia species using piecewise regression analyses. Finally, we evaluate how variation in the number of co-flowering Clarkia species in the community affects the strength of pollen quality and quality limitation. ResultsAcross all contexts, pollen deposition and the proportion of pollen tubes produced varied greatly among individuals, populations and species, and these were not always correlated. For instance, C. xantiana received the smallest pollen loads yet produced the highest proportion of pollen tubes, while C. speciosa exhibited the opposite pattern. Yet, co-flowering richness had variable effects on the strength of pollen quantity and quality limitation among populations. Specifically, breakpoint values, which are an indicator of overall pollen limitation, were two-fold higher in the four-species community compared with one- and two-species communities for two Clarkia species, suggesting that pollen limitation can increase with increasing richness of co-flowering congeners. ConclusionsOur results reveal a complex interplay between the quantity and quality of pollen limitation and co-flowering context that may have different evolutionary outcomes across species and populations. 

Abstract Determining how pollinators visit plants vs. how they carry and transfer pollen is an ongoing project in pollination ecology. The current tools for identifying the pollens that bees carry have different strengths and weaknesses when used for ecological inference. In this study we use three methods to better understand a system of congeneric, coflowering plants in the genusClarkiaand their bee pollinators: observations of plant–pollinator contact in the field, and two different molecular methods to estimate the relative abundance of eachClarkiapollen in samples collected from pollinators. We use these methods to investigate if observations of plant–pollinator contact in the field correspond to the pollen bees carry; if individual bees carryClarkiapollens in predictable ways, based on previous knowledge of their foraging behaviors; and how the three approaches differ for understanding plant–pollinator interactions. We find that observations of plant–pollinator contact are generally predictive of the pollens that bees carry while foraging, and network topologies using the three different methods are statistically indistinguishable from each other. Results from molecular pollen analysis also show that while bees can carry multiple species ofClarkiaat the same time, they often carry one species of pollen. Our work contributes to the growing body of literature aimed at resolving how pollinators use floral resources. We suggest our novel relative amplicon quantification method as another tool in the developing molecular ecology and pollination biology toolbox. 

Abstract Spatial partitioning is a classic hypothesis to explain plant species coexistence, but evidence linking local environmental variation to spatial sorting, demography and species' traits is sparse. If co‐occurring species' performance is optimized differently along environmental gradients because of trait variation, then spatial variation might facilitate coexistence.We used a system of four naturally co‐occurring species ofClarkia(Onagraceae) to ask whether distribution patchiness corresponds to variation in two environmental variables that contribute to hydrological variation. We then reciprocally sowedClarkiainto each patch type and measured demographic rates in the absence of congeneric competition. Species sorted in patches along one or both gradients, and in three of the four species, germination rate in the ‘home’ patch was higher than all other patches.Spatially variable germination resulted in the same three species exhibiting the highest population growth rates in their home patches.Species' trait values related to plant water use, as well as indicators of water stress in home patches, differed among species and corresponded to home patch attributes. However, post‐germination survival did not vary among species or between patch types, and fecundity did not vary spatially.Synthesis. Our research demonstrates the likelihood that within‐community spatial heterogeneity affects plant species coexistence, and presents novel evidence that differential performance in space is explained by what happens in the germination stage. Despite the seemingly obvious link between adult plant water‐use and variation in the environment, our results distinguish the germination stage as important for spatially variable population performance. 

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. 

Facilitation is likely important for understanding community diversity dynamics, but its myriad potential mechanisms are under-investigated. Studies of pollinator-mediated facilitation in plants, for example, are typically focused on how co-flowering species facilitate each other's pollination within a season. However, pollinator-mediated facilitation could also arise in the form of inter-annual pollination support, where co-occurring plant populations mutually facilitate each other by providing dynamic stability to support a pollinator population through time. In this work, I test this hypothesis with simulation models of annual flowering plant and bee pollinator populations to determine if and how inter-annual pollination support affects the persistence and/or stability of simulated communities. Two-species plant communities persisted at higher rates than single-species communities, and facilitation was strongest in communities with low mean germination rates and highly species-specific responses to environmental variation. Single-species communities were often more stable than their counterparts, likely because of survivorship—persistent single-species communities were necessarily more stable through time to support pollinators. This work shows that competition and facilitation can simultaneously affect plant population dynamics. It also importantly identifies key features of annual plant communities that might exhibit inter-annual pollination support- those with low germination rates and species-specific responses to variation. 

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, though delayed germination increased long-term fitness in 7 of 20 populations. Optimal germination fractions were 2 to 5 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 insufficient 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.