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Abstract Heritable symbionts are often observed at intermediate prevalence within host populations, despite expectations that positive fitness feedbacks should drive beneficial symbionts to fixation. Intermediate prevalence may reflect neutral dynamics of symbionts with weak fitness effects, transient dynamics of symbionts trending towards fixation (or elimination), or a stable intermediate outcome determined by the balance of fitness effects and failed symbiont transmission. Theory suggests that these outcomes should depend on symbiont‐conferred demographic effects and vertical transmission efficiency, which may both depend on environmental context.
We established experimental populations of winter bent grass
Agrostis hyemalis across a range of prevalence of the heritable fungal endophyteEpichloë amarillans . Using irrigation, we elevated the precipitation for half of the populations, which we hypothesized would weaken the benefits of symbiosis. Across two annual transitions, we assayed 5,485 individuals to determine prevalence and censused 954 individuals for demographic (survival, flowering, reproduction and recruitment) and vertical transmission data. We used hierarchical Bayesian models to infer long‐run equilibria from short‐term changes in symbiont prevalence and estimated demographic vital rates to link individual‐level effects to population‐level outcomes.We found evidence for all three proposed mechanisms for intermediate symbiont prevalence, but the outcome differed qualitatively across years and precipitation treatments. In the first year, symbionts trended towards fixation under drought conditions but drifted neutrally under elevated precipitation. Fixation likely arose from symbiont‐conferred recruitment benefits outweighing reproductive costs under the drought conditions, while elevated precipitation tempered these effects. In the second transition year, we inferred stable intermediate prevalence across both precipitation treatments, which indicated a balance between symbiont conferred recruitment benefits that allowed low‐prevalence populations to increase and imperfect transmission that caused high‐prevalence populations to decrease.
Synthesis . We find support for neutral, transient and stable mechanisms underlying symbiont prevalence, indicating that symbiont prevalence is often pushed and pulled in different directions by the composite outcome of symbiont effects on demographic rates and transmission efficiency, and the way in which these processes respond to environmental context. -
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Premise Variation in pollinator effectiveness may contribute to pollen limitation in fragmented plant populations. In plants with multiovulate ovaries, the number of conspecific pollen grains per stigma often predicts seed set and is used to quantify pollinator effectiveness. In the Asteraceae, however, florets are uniovulate, which suggests that the total amount of pollen deposited per floret may not measure pollinator effectiveness. We examined two aspects of pollinator effectiveness—effective pollen deposition and effective pollen movement—for insects visiting
Echinacea angustifolia , a composite that is pollen limited in small, isolated populations.Methods We filmed insect visits to
Echinacea in two prairie restorations and used these videos to quantify behavior that might predict effectiveness. To quantify effective pollen deposition, we used the number of styles shriveled per visit. To quantify effective pollen movement, we conducted paternity analysis on a subset of offspring and measured the pollen movement distance between mates.Results Effective pollen deposition varied among taxa.
Andrena helianthiformis , a Heliantheae oligolege, was the most effective taxon, shriveling more than twice the proportion of styles as all other visitors. Differences in visitor behavior on a flowering head did not explain variation in effective pollen deposition, nor did flowering phenology. On average, visitors moved pollen 16 m between plants, and this distance did not vary among taxa.Conclusions Andrena helianthiformis is an important pollinator ofEchinacea . Variation in reproductive fitness ofEchinacea in fragmented habitat may result, in part, from the abundance of this species.
