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  1. A long-standing problem in the study of mutualism is to understand the effects of non-mutualistic community members that exploit the benefits of mutualism without offering commodities in exchange (i.e., ‘exploiters’). Mutualisms are continually challenged by exploiters and their persistence may depend on the costliness of exploitation or on adaptations that allow mutualists to avoid the negative effects of exploiters. Coevolution could lead to changes in mutualists and exploiters that allow mutualisms to persist. Although coevolution is considered essential for mutualism persistence and resistance to disturbance, we have yet to obtain direct experimental evidence of the role of coevolution in resistance to exploitation. Additionally, resistance to exploitation via coevolutionary processes might vary with the degree of dependency between mutualistic partners, as facultative mutualisms are thought to be under weaker coevolutionary selection than obligate mutualisms. Here, we conducted an experimental evolution study using a synthetic yeast mutualism to test how coevolution in facultative and obligate mutualisms affects their resistance to exploitation. We found that naïve facultative mutualisms were more likely to breakdown under exploitation than naïve obligate mutualisms. After 15 weeks of coevolution, both facultative and obligate evolved mutualists were more likely to survive exploitation than naïve mutualists when we reassembled mutualist communities. Additionally, coevolved exploiters were more likely to survive with mutualists, whereas naïve exploiters frequently went extinct. These results suggest that coevolution between mutualists and exploiters can lead to mutualism persistence, potentially explaining why exploitation is ubiquitous but rarely associated with mutualism breakdown.

     
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    Free, publicly-accessible full text available March 14, 2025
  2. Abstract

    Functional traits fall along a continuum from resource conservative to acquisitive and are powerful predictors of the ecological settings necessary for a species to persist and establish. As a consequence, a major problem that functional trait analysis could address is understanding the ecological contexts necessary for the persistence of polyploid plants, because early generation polyploids, or “neopolyploids,” are at a high extinction risk. Because neopolyploidy could increase nutrient limitation, growth strategies should shift to accommodate the increased need for resources, but this prediction is untested. To address this gap, we compared the functional trait responses of diploids, synthetic neotetraploids, and naturally occurring tetraploids ofHeuchera cylindrica, an herbaceous perennial plant, to nutrient manipulations in a greenhouse experiment. We found strong support for the hypothesis that neotetraploidy increases nutrient requirements, as evidenced by reduced productivity and increased tissue concentrations of nitrogen and phosphorus in neotetraploids. We also found that the repeated formation of independent origins of neotetraploidy led to differing responses to nutrient supply, but neotetraploidy generally shifted functional traits to be more resource acquisitive and inefficient. Taken together, our results suggest that shifts in functional trait responses may constrain the ability of neopolyploids to establish in nutrient‐poor habitats.

     
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  3. Abstract

    Host plant shifts are central to diversification in insect herbivores. Many mechanisms can cause host shifts in insects, but one relatively unexplored mechanism is whole‐genome duplication (WGD) in the host plant. WGD, or polyploidy, is common in plants and causes spontaneous changes in physiology, morphology, and palatability that could impact the ability of herbivores to feed and develop on newly formed polyploids (neopolyploids).

    Here the authors tested if WGD affected the preference and performance of the specialist aphid,Acyrthosiphon pisum(pea aphids). Pea aphids seasonally form specialised lineages or ‘host forms’ on many host plant species including alfalfa and red clover. Aphid host forms on alfalfa and red clover naturally exist on different cytotypes of their respective hosts, with red clover aphids feeding on diploid clover and alfalfa aphids feeding on tetraploid alfalfa. Therefore, the authors predicted that these host forms would have a higher preference for and performance on their respective natal host cytotype.

    Neither host form exhibited a preference for a particular cytotype, but there were modest changes in aphid performance based on host cytotype. Specifically, aphids specialised to red clover had higher fecundity on diploid red clover than on neotetraploid red clover. Together, these results showed that both host forms were able to recognise and accept different cytotypes of the two host species, but only one host form experienced trade‐offs in performance when feeding on neotetraploids. These results suggest that WGD may act as a mechanism of host expansion in pea aphids as plants speciate via WGD.

     
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  5. Abstract

    Species interactions shape the evolution of traits, life histories and the pattern of speciation. What is less clear is whether certain types of species interaction are more or less likely to lead to phenotypic divergence among species. We used the brood pollination mutualism between yuccas and yucca moths to test how mutualistic (pollination) and antagonistic (oviposition) traits differ in the propensity to increase phenotypic divergence among pollinator moths. We measured traits of the tentacular mouthparts, structures used by females to actively pollinate flowers, as well as ovipositor traits to examine differences in the rate of evolution of these two suites of traits among pollinator species. Morphological analyses revealed two distinct groups of moths based on ovipositor morphology, but no such groupings were identified for tentacle morphology, even for moths that pollinated distantly related yuccas. In addition, ovipositor traits evolved at significantly faster rates than tentacular traits. These results support theoretical work suggesting that antagonism is more likely than mutualism to lead to phenotypic divergence.

     
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  6. Premise

    Polyploidy is known to cause physiological changes in plants which, in turn, can affect species interactions. One major physiological change predicted in polyploid plants is a heightened demand for growth‐limiting nutrients. Consequently, we expect polyploidy to cause an increased reliance on the belowground mutualists that supply these growth‐limiting nutrients. An important first step in investigating how polyploidy affects nutritional mutualisms in plants, then, is to characterize differences in the rate at which diploids and polyploids interact with belowground mutualists.

    Methods

    We usedHeuchera cylindrica(Saxifragaceae) to test how polyploidy influences interactions with arbuscular mycorrhizal fungi (AMF). Here we first confirmed the presence ofAMFinH. cylindrica, and then we used field‐collected specimens to quantify and compare the presence ofAMFstructures while controlling for site‐specific variation.

    Results

    Tetraploids had higher colonization rates as measured by total, hyphal, and nutritional‐exchange structures; however, we found that diploids and tetraploids did not differ in vesicle colonization rates.

    Conclusions

    The results suggest that polyploidy may alter belowground nutritional mutualisms with plants. Because colonization by nutritional‐exchange structures was higher in polyploids but vesicle colonization was not, polyploids might form stronger associations with theirAMFpartners. Controlled experiments are necessary to test whether this pattern is driven by the direct effect of polyploidy onAMFcolonization.

     
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  7. Premise

    Although polyploidy has been studied since the early 1900s, fundamental aspects of polyploid ecology and evolution remain unexplored. In particular, surprisingly little is known about how newly formed polyploids (neopolyploids) become demographically established. Models predict that most polyploids should go extinct within the first few generations as a result of reproductive disadvantages associated with being the minority in a primarily diploid population (i.e., the minority cytotype principle), yet polyploidy is extremely common. Therefore, a key goal in the study of polyploidy is to determine the mechanisms that promote polyploid establishment in nature. Because premating isolation is critical in order for neopolylpoids to avoid minority cytotype exclusion and thus facilitate establishment, we examined floral morphology and three common premating barriers to determine their importance in generating reproductive isolation of neopolyploids from diploids.

    Methods

    We induced neopolyploidy inTrifolium pratenseand compared their floral traits to the diploid progenitors. In addition to shifts in floral morphology, we examined three premating barriers: isolation by self‐fertilization, flowering‐time asynchrony, and pollinator‐mediated isolation.

    Results

    We found significant differences in the morphology of diploid and neopolyploid flowers, but these changes did not facilitate premating barriers that would generate reproductive isolation of neopolyploids from diploids. There was no difference in flowering phenology, pollinator visitation, or selfing between the cytotypes.

    Conclusions

    Our results indicate that barriers other than the ones tested in this study—such as geographic isolation, vegetative reproduction, and pistil–stigma incompatibilities—may be more important in facilitating isolation and establishment of neopolyploidT. pratense.

     
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