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  1. Premise

    Whether drought‐adaptation mechanisms tend to evolve together, evolve independently, or evolve constrained by genetic architecture is incompletely resolved, particularly for water‐relations traits besides gas exchange. We addressed this issue in two subspecies ofClarkia xantiana(Onagraceae), California winter annuals that separated approximately 65,000 years ago and are adapted, partly by differences in flowering time, to native ranges differing in precipitation.


    In these subspecies and in recombinant inbred lines (RILs) from a cross between them, we scored traits related to drought adaptation (timing of seed germination and of flowering, succulence, pressure–volume curve variables) in common environments.


    The subspecies native to more arid environments (parviflora) exhibited slower seed germination in saturated conditions, earlier flowering, and greater succulence, likely indicating superior drought avoidance, drought escape, and dehydration resistance via water storage. The other subspecies (xantiana) had lower osmotic potential at full turgor and lower water potential at turgor loss, implying superior dehydration tolerance. Genetic correlations among RILs suggest facilitated evolution of some trait combinations and independence of others. Where genetic correlations exist, subspecies differences fell along them, with the exception of differences in succulence and turgor loss point. In that case, subspecies difference overcame genetic correlations, possibly reflecting strong selection and/or antagonistic genetic correlations with other traits.


    Clarkia xantianasubspecies’ differ in multiple mechanisms of drought adaptation. Genetic architecture generally does not seem to have constrained the evolution of these mechanisms, and it may have facilitated the evolution of some of trait combinations.

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  2. 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.

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