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Thompson, Ken A.; Renaudin, Marie; Johnson, Marc T.J. 2016-11-16 Urban ecosystems are an increasingly dominant feature of terrestrial landscapes. While evidence that species can adapt to urban environments is accumulating, the mechanisms through which urbanization imposes natural selection on populations are poorly understood. The identification of adaptive phenotypic changes (i.e. clines) along urbanization gradients would facilitate our understanding of the selective factors driving adaptation in cities. Here, we test for phenotypic clines in urban ecosystems by sampling the frequency of a Mendelian-inherited trait—cyanogenesis—in white clover (Trifolium repens L.) populations along urbanization gradients in four cities. Cyanogenesis protects plants from herbivores, but reduces tolerance to freezing temperatures. We found that the frequency of cyanogenic plants within populations decreased towards the urban centre in three of four cities. A field experiment indicated that spatial variation in herbivory is unlikely to explain these clines. Rather, colder minimum winter ground temperatures in urban areas compared with non-urban areas, caused by reduced snow cover in cities, may select against cyanogenesis. In the city with no cline, high snow cover might protect plants from freezing damage in the city centre. Our study suggests that populations are adapting to urbanization gradients, but regional climatic patterns may ultimately determine whether adaptation occurs.
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Thompson, Ken A.; Husband, Brian C.; Maherali, Hafiz 2015-09-23 Premise of the study: Polyploidy—the possession of more than two copies of each chromosome in the nucleus—is common in flowering plants. Polyploid plants can occupy different geographic ranges than their diploid progenitors, but the factors responsible for maintaining these range differences are poorly understood. Polyploidy can have significant physiological consequences, and the present study aims to determine whether previously described physiological differences between cytotypes are correlated with climatic niches and geographic distributions. Methods: Prior research indicates that tetraploid Chamerion angustifolium (Onagraceae) are more tolerant of drought and less tolerant of freezing than diploids, which suggests that they should occupy a niche that is warmer and drier than that of diploids. We extracted climate data for 134 C. angustifolium populations classified as pure diploid, pure tetraploid, or mixed-ploidy. We compared climatic conditions between these population categories, and generated ecological niche models to compare their geographic distribution with prior qualitative estimates. Key results: Pure tetraploid populations occupy habitats that are warmer and drier than pure diploid populations. Mixed-ploidy populations occur in habitats that are not strictly intermediate between pure diploid and pure tetraploid populations, but are as cold as pure diploid populations and have intermediate soil moisture deficits. Our niche models were similar to previous qualitative estimates of cytotype geographic distribution. Conclusions: The correspondence between the physiological tolerances of cytotypes, their climatic niches, and their geographic distributions suggests that physiological traits are at least partially responsible for differences in the realized climatic niches of diploid and tetraploid C. angustifolium. https://creativecommons.org/publicdomain/zero/1.0/
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Thompson, Kenneth A.; Johnson, Marc T. J.; Thompson, Ken A. 2016-03-21 While many studies demonstrate that herbivores alter selection on plant reproductive traits, little is known about whether antiherbivore defenses affect selection on these traits. We hypothesized that antiherbivore defenses could alter selection on reproductive traits by altering trait expression through allocation trade-offs, or by altering interactions with mutualists and/or antagonists. To test our hypothesis, we used white clover, Trifolium repens, which has a Mendelian polymorphism for the production of hydrogen cyanide—a potent antiherbivore defense. We conducted a common garden experiment with 185 clonal families of T. repens that included cyanogenic and acyanogenic genotypes. We quantified resistance to herbivores, and selection on 6 floral traits and phenology via male and female fitness. Cyanogenesis reduced herbivory but did not alter the expression of reproductive traits through allocation trade-offs. However, the presence of cyanogenic defenses altered natural selection on petal morphology and the number of flowers within inflorescences via female fitness. Herbivory influenced selection on flowers and phenology via female fitness independently of cyanogenesis. Our results demonstrate that both herbivory and antiherbivore defenses alter natural selection on plant reproductive traits. We discuss the significance of these results for understanding how antiherbivore defenses interact with herbivores and pollinators to shape floral evolution.
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Thompson, Kenneth A.; Husband, Brian C.; Maherali, Hafiz; Thompson, Ken A. 2016-02-09 1. Polyploid plants often occupy different geographic ranges than their diploid progenitors, but the causes of this segregation are poorly understood. Differential competitive abilities of cytotypes across an environmental gradient could be responsible for these observed geographic range differences. 2. Cytotypes of Chamerion angustifolium (Onagraceae) are mostly allopatric, and prior research indicates that tetraploids are more physiologically tolerant of water limitation and occupy drier habitats than diploids. We hypothesized that tetraploids are stronger competitors than diploids in soils where water is limited, which allows them to persist in dry habitats while diploids cannot. 3. We grew both cytotypes together in competition under water-limited and well-watered conditions. We varied both total plant density and the relative frequency of cytotypes among pots, which allowed us to separate the effects of intra-cytotypic and inter-cytotypic competition. 4. Both diploid and tetraploid plants were smaller in the water-limited treatment than in the well-watered treatment. Nevertheless, there were no differences in the relative strength of intra-cytotypic and inter-cytotypic competition experienced by either cytotype across the watering treatments, indicating that diploids and tetraploids had equal competitive abilities in both treatments. 5. Synthesis. Competition for limiting resources is often proposed as a mechanism causing ecological and geographic segregation between diploid and polyploid cytotypes. Our results do not support the hypothesis that tetraploid Chamerion angustifolium plants are stronger competitors than diploids when water is limited. A differential ability to compete for water is likely not responsible for the observed ecological and geographic segregation between cytotypes in this species. Competition may not be a general mechanism that causes segregation between diploid and polyploid cytotypes in nature.
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Santangelo, James S.; Thompson, Ken A.; Johnson, Marc T. J. 2020-03-20 Pollinators and herbivores can both affect the evolutionary diversification of plant reproductive traits. However, plant defenses frequently alter antagonistic and mutualistic interactions and therefore variation in plant defenses may alter patterns of herbivore- and pollinator-mediated selection on plant traits. We tested this hypothesis by conducting a common garden field experiment using 50 clonal genotypes of white clover (Trifolium repens) that varied in a Mendelian inherited chemical antiherbivore defense—the production of hydrogen cyanide (HCN). To evaluate whether plant defenses alter herbivore- and/or pollinator-mediated selection, we factorially crossed chemical defense (25 cyanogenic and 25 acyanogenic genotypes), herbivore damage (herbivore suppression) and pollination (hand-pollination). We found that herbivores weakened selection for increased inflorescence production, suggesting that large displays are costly in the presence of herbivores. In addition, herbivores weakened selection on flower size but only among acyanogenic plants, suggesting that plant defenses reduce the strength of herbivore-mediated selection. Pollinators did not independently affect selection on any trait, although pollinators weakened selection for later flowering among cyanogenic plants. Overall, cyanogenic plant defenses consistently increased the strength of positive directional selection on reproductive traits. Herbivores and pollinators both strengthened and weakened the strength of selection on reproductive traits, although herbivores imposed ~2.7× stronger selection than pollinators across all traits. Contrary to the view that pollinators are the most important agents of selection on reproductive traits, our data show that selection on reproductive traits is driven primarily by variation in herbivory and plant defenses in this system.

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