Among-population variation in tolerance to larval herbivory by Anthocharis cardamines in the polyploid herb Cardamine pratensis.

Plants have two principal defense mechanisms to decrease fitness losses to herbivory: tolerance, the ability to compensate fitness after damage, and resistance, the ability to avoid damage. Variation in intensity of herbivory among populations should result in variation in plant defense levels if tolerance and resistance are associated with costs. Yet little is known about how levels of tolerance are related to resistance and attack intensity in the field, and about the costs of tolerance. In this study, we used information about tolerance and resistance against larval herbivory by the butterfly Anthocharis cardamines under controlled conditions together with information about damage in the field for a large set of populations of the perennial plant Cardamine pratensis. Plant tolerance was estimated in a common garden experiment where plants were subjected to a combination of larval herbivory and clipping. We found no evidence of that the proportion of damage that was caused by larval feeding vs. clipping influenced plant responses. Damage treatments had a negative effect on the three measured fitness components and also resulted in an earlier flowering in the year after the attack. Tolerance was related to attack intensity in the population of origin, i.e. plants from populations with higher attack intensity were more likely to flower in the year following damage. However, we found no evidence of a relationship between tolerance and resistance. These results indicate that herbivory drives the evolution for increased tolerance, and that changes in tolerance are not linked to changes in resistance. We suggest that the simultaneous study of tolerance, attack intensity in the field and resistance constitutes a powerful tool to understand how plant strategies to avoid negative effects of herbivore damage evolve.

Novel antagonistic interactions associated with plant polyploidization influence trait selection and habitat preference.

Polyploidization is an important mechanism for sympatric speciation in plants. Still, we know little about whether plant polyploidization leads to insect host shifts, and if novel interactions influence habitat and trait selection in plants. We investigated herbivory by the flower bud gall-forming midge Dasineura cardaminis on tetraploids and octoploids of the herb Cardamine pratensis. Gall midges attacked only octoploid plant populations, and a transplantation experiment confirmed this preference. Attack rates were higher in populations that were shaded, highly connected or occurred along stream margins. Within populations, late-flowering individuals with many flowers were most attacked. Galling reduced seed production and significantly influenced phenotypic selection on flower number. Our results suggest that an increase in ploidy may lead to insect host shifts and that plant ploidy explains insect host use. In newly formed plant polyploids, novel interactions may alter habitat preferences and trait selection, and influence the further evolution of cytotypes.

Butterfly seed predation: effects of landscape characteristics, plant ploidy level and population structure.

Polyploidization has been suggested as one of the most common mechanisms for plant diversification. It is often associated with changes in several morphological, phenological and ecological plant traits, and therefore has the potential to alter insect-plant interactions. Nevertheless, studies evaluating the effect of plant polyploidy on interspecific interactions are still few. We investigated pre-dispersal seed predation by the butterfly Anthocharis cardamines in 195 populations of two ploidy levels of the herb Cardamine pratensis (tetraploid ssp. pratensis, 2n = 30 vs. octoploid ssp. paludosa, 2n = 56-64). We asked if differences in incidence and intensity of predation among populations were related to landscape characteristics, plant ploidy level and population structure. The incidence of the seed predator increased with increasing plant population size and decreasing distance to nearest population occupied by A. cardamines. The intensity of predation decreased with increasing plant population size and was not affected by isolation. Probability of attack decreased with increasing shading, and intensity of predation was higher in grazed than in non-grazed habitats. The attack intensity increased with increasing mean flower number of plant population, but was not affected by flowering phenology. Individuals in tetraploid populations suffered on average from higher levels of seed predation, had higher mean flower number, were less shaded and occurred more often in grazed habitats than octoploid populations. When accounting for differences in habitat preferences between ploidy levels there was no longer a difference in intensity of predation, suggesting that the observed differences in attack rates among populations of the two ploidy levels are mediated by the habitat. Overall, our results suggest that polyploidization is associated with differentiation in habitat preferences and phenotypic traits leading to differences in interspecific interaction among plant populations. This, in turn, may facilitate further divergence of ploidy levels.