word count: 237 words 19 Total word count: 5095 words 20 Abstract 21Genetic variation for partner quality in mutualisms is an evolutionary paradox. One possible 22 resolution to this puzzle is that there is a tradeoff between partner quality and other fitness-23 related traits. Here, we tested whether a susceptibility to parasitism is one such tradeoff in the 24 mutualism between legumes and nitrogen-fixing bacteria (rhizobia). We performed two 25 greenhouse experiments with the legume Medicago truncatula. In the first, we inoculated each 26 plant with the rhizobia Ensifer meliloti and with one of 40 genotypes of the parasitic root-knot 27 nematode Meloidogyne hapla. In the second experiment, we inoculated all plants with rhizobia 28 and half of the plants with a genetically variable population of nematodes. Using the number of 29 nematode galls as a proxy for infection severity, we found that plant genotypes differed in 30 susceptibility to nematode infection, and nematode genotypes differed in infectivity. Second, we 31 showed that there was a genetic correlation between the number of mutualistic structures formed 32 by rhizobia (nodules) and the number of parasitic structures formed by nematodes (galls). 33Finally, we found that nematodes disrupt the rhizobia mutualism: nematode-infected plants 34 formed fewer nodules and had less nodule biomass than uninfected plants. Our results 35 demonstrate that there is genetic conflict between attracting rhizobia and repelling nematodes in 36Medicago. If genetic conflict with parasitism is a general feature of mutualism, it could account 37 for the maintenance of genetic variation in partner quality and influence the evolutionary 38 dynamics of positive species interactions. 39 40 Impact summary 41Cooperative species interactions, known as mutualisms, are vital for organisms from plants to 42 humans. For example, beneficial microbes in the human gut are a necessary component of 43 digestive health. However, parasites often infect their hosts via mechanisms that are 44 extraordinarily similar to those used by mutualists, which may create a tradeoff between 45 attracting mutualists and resisting parasites. In this study, we investigated whether this tradeoff 46 exists, and how parasites impact mutualism function in the barrelclover Medicago truncatula, a 47 close relative of alfalfa. Legumes like Medicago depend on nitrogen provided by mutualistic 48 bacteria (rhizobia) to grow, but they are also infected by parasitic worms called nematodes, 49 which steal plant nutrients. Both microorganisms live in unique structures (nodules and galls) on 50 plant roots. We showed that the benefits of mutualism and the costs of parasitism are predicted 51 by the number of mutualistic structures (nodules) and the number of parasitic structures (galls), 52 respectively. Second, we found that there is a genetic tradeoff between attracting mutualists and 53 repelling parasites in Medicago truncatula: plant genotypes that formed more rhizobia nodules 54 also formed more nematode galls. Final...

Intraspecific variation in flower color is often attributed to pollinator-mediated selection, yet this mechanism cannot explain flower color polymorphisms in self-pollinating species. Indirect selection mediated via biotic and abiotic stresses could maintain flower color variation in these systems. The selfing forb, Boechera stricta, typically displays white flowers, but some individuals produce purple flowers. We quantified environmental correlates of flower color in natural populations. To disentangle plasticity from genotypic variation, we performed a multiyear field experiment in five gardens. In controlled conditions, we evaluated herbivore preferences and the effects of drought stress and soil pH on flower color expression. In natural populations, purple-flowered individuals experienced lower foliar herbivory than did their white-flowered counterparts. This pattern also held in the common gardens. Additionally, low-elevation environments induced pigmented flowers (plasticity), and the likelihood of floral pigmentation decreased with source elevation of maternal families (genetic cline). Viability selection favored families with pigmented flowers. In the laboratory, herbivores exerted greater damage on tissue derived from white- vs purple-flowered individuals. Furthermore, drought induced pigmentation in white-flowered lineages, and white-flowered plants had a fecundity advantage in the well-watered control. Flower color variation in selfing species is probably maintained by herbivory, drought stress, and other abiotic factors that vary spatially.

Genetic variation for partner quality in mutualisms is an evolutionary paradox. One possible resolution to this puzzle is that there is a tradeoff between partner quality and other fitness‐related traits. Here, we tested whether susceptibility to parasitism is one such tradeoff in the mutualism between legumes and nitrogen‐fixing bacteria (rhizobia). We performed two greenhouse experiments with the legume Medicago truncatula. In the first, we inoculated each plant with the rhizobia Ensifer meliloti and with one of 40 genotypes of the parasitic root‐knot nematode Meloidogyne hapla. In the second experiment, we inoculated all plants with rhizobia and half of the plants with a genetically variable population of nematodes. Using the number of nematode galls as a proxy for infection severity, we found that plant genotypes differed in susceptibility to nematode infection, and nematode genotypes differed in infectivity. Second, we showed that there was a genetic correlation between the number of mutualistic structures formed by rhizobia (nodules) and the number of parasitic structures formed by nematodes (galls). Finally, we found that nematodes disrupt the rhizobia mutualism: nematode‐infected plants formed fewer nodules and had less nodule biomass than uninfected plants. Our results demonstrate that there is genetic conflict between attracting rhizobia and repelling nematodes in Medicago. If genetic conflict with parasitism is a general feature of mutualism, it could account for the maintenance of genetic variation in partner quality and influence the evolutionary dynamics of positive species interactions.

The maintenance of genetic variation in mutualism-related traits is key for understanding mutualism evolution, yet the mechanisms maintaining variation remain unclear. We asked whether genotype-by-environment (G×E) interaction is a potential mechanism maintaining variation in the model legume–rhizobia system, Medicago truncatula–Ensifer meliloti . We planted 50 legume genotypes in a greenhouse under ambient light and shade to reflect reduced carbon availability for plants. We found an expected reduction under shaded conditions for plant performance traits, such as leaf number, aboveground and belowground biomass, and a mutualism-related trait, nodule number. We also found G×E for nodule number, with ∼83% of this interaction due to shifts in genotype fitness rank order across light environments, coupled with strong positive directional selection on nodule number regardless of light environment. Our results suggest that G×E can maintain genetic variation in a mutualism-related trait that is under consistent positive directional selection across light environments.