A direct comparison of the consequences of plant genotypic and species diversity on communities and ecosystem function

Biodiversity loss is proceeding at an unprecedented rate, yet we lack a thorough understanding of the consequences of losing diversity at different scales. While species diversity is known to impact community and ecosystem processes, genotypic diversity is assumed to have relatively smaller effects. Nonetheless, a few recent studies suggest that genotypic diversity may have quantitatively similar ecological consequences compared to species diversity. Here we show that increasing either genotypic diversity of common evening primrose (Oenothera biennis) or species diversity of old?field plant species resulted in nearly equivalent increases (?17%) in aboveground primary production. The predominant mechanism explaining this effect, niche complementarity, was similar for each type of diversity. Arthropod species richness also increased with both types of plant diversity, but the mechanisms leading to this effect differed: abundance?driven accumulation of arthropod species was important in plant genotypic polycultures, whereas resource specialization was important in plant species polycultures. Thus, similar increases in primary productivity differentially impacted higher trophic levels in response to each type of plant diversity. These results highlight important ecological similarities and differences between genotypic and species diversity and suggest that genotypic diversity may play a larger role in community and ecosystem processes than previously realized.The NSF (DEB- 0950231 to A. A. Agrawal and IGERT small grant in Biogeochemistry and Environmental Biocomplexity to S. H. McArt, S. C. Cook-Patton, and A. L. Parachnowitsch) and USDA-NRI (2006-35302-17431 to J. S. Thaler) supported this wor

Pollinators and herbivores interactively shape selection on strawberry defence and attraction

Tripartite interactions between plants, herbivores, and pollinators hold fitness consequences for most angiosperms. However, little is known on how plants evolve in response-and in particular what the net selective outcomes are for traits of shared relevance to pollinators and herbivores. In this study, we manipulated herbivory ("presence" and "absence" treatments) and pollination ("open" and "hand pollination" treatments) in a full factorial common-garden experiment with woodland strawberry (Fragaria vesca L.). This design allowed us to quantify the relative importance and interactive effects of herbivore- and pollinator-mediated selection on nine traits related to plant defence and attraction. Our results showed that pollinators imposed stronger selection than herbivores on traits related to both direct and indirect (i.e., tritrophic) defence. However, conflicting selection was imposed on inflorescence density: a trait that appears to be shared by herbivores and pollinators as a host plant signal. However, in all cases, selection imposed by one agent depended largely on the presence or ecological effect of the other, suggesting that dynamic patterns of selection could be a common outcome of these interactions in natural populations. As a whole, our findings highlight the significance of plant-herbivore-pollinator interactions as potential drivers of evolutionary change, and reveal that pollinators likely play an underappreciated role as selective agents on direct and in direct plant defence.</p

Measuring, comparing and interpreting phenotypic selection on floral scent

Natural selection on floral scent composition is a key element of the hypothesis that pollinators and other floral visitors drive scent evolution. The measure of such selection is complicated by the high-dimensional nature of floral scent data and uncertainty about the cognitive processes involved in scent-mediated communication. We use dimension reduction through reduced-rank regression to jointly estimate a scent composite trait under selection and the strength of selection acting on this trait. To assess and compare variation in selection on scent across species, time and space, we reanalyse 22 datasets on six species from four previous studies. The results agreed qualitatively with previous analyses in terms of identifying populations and scent compounds subject to stronger selection but also allowed us to evaluate and compare the strength of selection on scent across studies. Doing so revealed that selection on floral scent was highly variable, and overall about as common and as strong as selection on other phenotypic traits involved in pollinator attraction or pollen transfer. These results are consistent with an important role of floral scent in pollinator attraction. Our approach should be useful for further studies of plant-animal communication and for studies of selection on other high-dimensional phenotypes. In particular, our approach will be useful for studies of pollinator-mediated selection on complex scent blends comprising many volatiles, and when no prior information on the physiological responses of pollinators to scent compounds is available

Measuring, comparing and interpreting phenotypic selection on floral scent

Natural selection on floral scent composition is a key element of the hypothesis that pollinators and other floral visitors drive scent evolution. The measure of such selection is complicated by the high-dimensional nature of floral scent data and uncertainty about the cognitive processes involved in scent-mediated communication. We use dimension reduction through reduced-rank regression to jointly estimate a scent composite trait under selection and the strength of selection acting on this trait. To assess and compare variation in selection on scent across species, time and space, we reanalyse 22 datasets on six species from four previous studies. The results agreed qualitatively with previous analyses in terms of identifying populations and scent compounds subject to stronger selection but also allowed us to evaluate and compare the strength of selection on scent across studies. Doing so revealed that selection on floral scent was highly variable, and overall about as common and as strong as selection on other phenotypic traits involved in pollinator attraction or pollen transfer. These results are consistent with an important role of floral scent in pollinator attraction. Our approach should be useful for further studies of plant-animal communication and for studies of selection on other high-dimensional phenotypes. In particular, our approach will be useful for studies of pollinator-mediated selection on complex scent blends comprising many volatiles, and when no prior information on the physiological responses of pollinators to scent compounds is available.Peer reviewe

Global urban environmental change drives adaptation in white clover

Urbanization transforms environments in ways that alter biological evolution. We examined whether urban environmental change drives parallel evolution by sampling 110,019 white clover plants from 6169 populations in 160 cities globally. Plants were assayed for a Mendelian antiherbivore defense that also affects tolerance to abiotic stressors. Urban-rural gradients were associated with the evolution of clines in defense in 47% of cities throughout the world. Variation in the strength of clines was explained by environmental changes in drought stress and vegetation cover that varied among cities. Sequencing 2074 genomes from 26 cities revealed that the evolution of urban-rural clines was best explained by adaptive evolution, but the degree of parallel adaptation varied among cities. Our results demonstrate that urbanization leads to adaptation at a global scale

Understanding Natural Selection On Floral Traits: Variation, Agents And Consequences

For animal pollinated plants, flowers are highly interactive organs expected to be under strong and potentially diverse selection pressures. Plants not only need to attract their pollinators but also deter their enemies, as well as keep the costs of these structures low. Thus natural selection on floral characters is an important area of inquiry. The following series of studies examines multiple aspects of natural selection on plants with a focus on floral characters. The first paper asks whether natural selection is more variable in space or time for both a focal species (Penstemon digitalis) and for in plants in general. Indeed, selection is variable, but not more so in either of these dimensions, suggesting that constraints to local adaptation and speciation are equally as likely as their occurrence. Knowing the agents of selection on traits can inform the direction and expectations for evolutionary change. The second paper discusses the finding that pollinators were driving selection for larger flowers and displays in P. digitalis. Furthermore, selection on floral traits was generally stronger when pollinators are present than when they were excluded for multiple species for which there is data. These results suggest that pollinators may indeed be major agents of natural selection on flowers. However, with so few studies directly testing agents of selection, this assumption should be applied with caution in contemporary populations. The third paper looks at an understudied trait in an evolutionary context. Although scents have been characterized for many plant species, very few studies examine the variation in this trait or how natural selection acts on it. Scents vary among populations of P. digitalis on a small geographic scale and there was significant natural selection to produce more scent in a common garden. Finally, the last paper examines the expected outcomes of selection for later flowering in Lobelia siphilitica by pre-dispersal seed predators. As predicted by optimal defence theory, flowering time and latex are correlated in this system suggesting that selection on flowering time could also effect selection on defence

Pollinators and herbivores interactively shape selection on strawberry defence and attraction

Tripartite interactions between plants, herbivores, and pollinators hold fitness consequences for most angiosperms. However, little is known on how plants evolve in response-and in particular what the net selective outcomes are for traits of shared relevance to pollinators and herbivores. In this study, we manipulated herbivory ("presence" and "absence" treatments) and pollination ("open" and "hand pollination" treatments) in a full factorial common-garden experiment with woodland strawberry (Fragaria vesca L.). This design allowed us to quantify the relative importance and interactive effects of herbivore- and pollinator-mediated selection on nine traits related to plant defence and attraction. Our results showed that pollinators imposed stronger selection than herbivores on traits related to both direct and indirect (i.e., tritrophic) defence. However, conflicting selection was imposed on inflorescence density: a trait that appears to be shared by herbivores and pollinators as a host plant signal. However, in all cases, selection imposed by one agent depended largely on the presence or ecological effect of the other, suggesting that dynamic patterns of selection could be a common outcome of these interactions in natural populations. As a whole, our findings highlight the significance of plant-herbivore-pollinator interactions as potential drivers of evolutionary change, and reveal that pollinators likely play an underappreciated role as selective agents on direct and in direct plant defence

Pollinator-mediated natural selection in Penstemon digitalis

Measuring the agents of natural selection is important because it allows us to understand not only which traits are expected to evolve but also why they will evolve. Natural selection by pollinators on floral traits is often assumed because in outcrossing animal-pollinated plants flowers are generally thought to function as advertisements of rewards directed at pollinators. We tested the role of bee pollinators in selection on Penstemon digitalis and found that pollinators were driving selection for larger and more flowers. However, what makes our publication unique is the additional information we gained from reviewing the few other studies that also directly tested whether pollinators were agents of selection on floral traits. As we would expect if pollinators are important agents of selection, selection on floral traits was significantly stronger when pollinators were present than when their choices were experimentally removed. Taken together, these results suggest that pollinators can be important drivers of selection in contemporary populations

Spatial variation in scent emission within flowers

Floral scent is considered an important long-distance signal that attracts pollinators, but also has been suggested to function at shorter distances such as within-flower nectar guides or as a defense against antagonists. Indeed, in some species floral scent production and emission show spatial patterns of variation within flowers, as certain compounds are exclusively emitted from specific floral tissues. In other species, the different volatile organic compounds that constitute the floral bouquet are emitted evenly from the entire flower. Here, we summarize the current evidence on floral scent variation within flowers by combining a literature review of published data on tissue-level floral scent variation (41 species) with floral scent dissections (17 species). For each species, we recorded the total number of volatile compounds separately and grouped in major chemical classes. To facilitate comparisons across diverse species, we compared volatiles emitted by 1) the whole flower, 2) the visual floral tissues (i.e. petals and colored structures), 3) non-visual floral tissues (i.e. green parts and reproductive structures), as well as 4) the compounds emitted by both visual and non-visual tissues. Results show that floral scent variation is frequent, but by no means ubiquitous, occurring in species from distantly related groups. We discuss the two main functional hypotheses promoting floral scent variation within flowers, i.e. as a pollinator attractant at short-distances or a defensive function against antagonists, together with non-functional hypotheses (e.g. pleiotropic effects, ecological costs). We point out further directions on this topic and suggest experimental approaches testing the attractiveness of compounds emitted by different floral parts alone and in combination with other floral signals. Our synthesis provides a foundation for future studies on the functional ecology of floral scent and reinforces the idea of high complexity in floral chemical signals

Differences in Floral Scent and Petal Reflectance Between Diploid and Tetraploid Chamerion angustifolium

Genome duplication in plants is thought to be a route to speciation due to cytotype incompatibility. However, to reduce cross-pollination between cytotypes in animal-pollinated species, distinctive floral phenotypes, which would allow pollinator-mediated assortative mating between flowers, are also expected. Chamerion angustifolium is a Holarctic species that forms a hybrid zone between diploid and tetraploid populations in the North American Rocky Mountains. Extensive research has shown that these cytotypes differ in many ways, including some floral traits, and that pollinators can discriminate between cytotypes, leading to assortative mating. However, two signals commonly used by insect pollinators have not been measured for this species, namely petal colour and floral scent. Using greenhouse-grown diploids and tetraploids of C. angustifolium from the ploidy hybrid-zone in the North American Rocky Mountains, we show that both floral scent signals and petal reflectance differ between cytotypes. These differences, along with differences in flower size shown previously, could help explain pollinator-mediated assortative mating observed in previous studies. However, these differences in floral phenotypes may vary in importance to pollinators. While the differences in scent included common floral volatiles readily detected by bumblebees, the differences in petal reflectance may not be perceived by bees based on their visual sensitivity across the spectra. Thus, our results suggest that differences in floral volatile emissions are more likely to contribute to pollinator discrimination between cytotypes and highlight the importance of understanding the sensory systems of pollinators when examining floral signals