No abstract

Greater plant defence is predicted to evolve at lower latitudes in response to increased herbivore pressure. However, recent studies question the generality of this pattern. In this study, we tested for genetically based latitudinal clines in resistance to herbivores and underlying defence traits of Oenothera biennis. We grew plants from 137 populations from across the entire native range of O. biennis. Populations from lower latitudes showed greater resistance to multiple specialist and generalist herbivores. These patterns were associated with an increase in total phenolics at lower latitudes. A significant proportion of the phenolics were driven by the concentrations of two major ellagitannins, which exhibited opposing latitudinal clines. Our analyses suggest that these findings are unlikely to be explained by local adaptation of herbivore populations or genetic variation in phenology. Rather greater herbivory at high latitudes can be explained by latitudinal clines in the evolution of plant defences.

The latitudinal herbivory-defense hypothesis (LHDH) posits that herbivory and plant defenses increase toward lower latitudes. Recent studies provide contradictory evidence and suggest alternative explanations for biogeographic patterns in plant-herbivore interactions. Here we test the LHDH by sampling herbivory from multiple generalist and specialist insect herbivores over the entire latitudinal native range of the plant species Oenothera biennis L. (Onagraceae). We sampled 79 populations on a 168 north-south gradient from Ontario and Maine to Alabama and Florida. From each population, we quantified herbivory across feeding guilds by considering leaf herbivory caused by generalist insects, damage by a specialist stem-boring beetle, and flower/seed herbivory by three Lepidoptera that specialize on Oenothera. We also related environmental and population density variables to herbivory. Our results show that latitudinal patterns vary dramatically among herbivore species. While generalist leaf herbivory showed no latitudinal pattern, stem borer damage increased with decreasing latitude. By contrast, the specialist flower/seed herbivores all caused less damage at lower latitudes. Temperature explained slightly more variation in herbivory than latitude, while precipitation and population density were less important. Overall, we show that every pattern of herbivory (positive, negative and no relationship) is possible across a latitudinal gradient, and this variation depends on the insects' degree of specialization and feeding guild.

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.

Uncovering the genetic basis of adaptation hinges on the ability to detect loci under selection. However, population genomics outlier approaches to detect selected loci may be inappropriate for clinal populations or those with unclear population structure because they require that individuals be clustered into populations. An alternate approach, landscape genomics, uses individualbased approaches to detect loci under selection and reveal potential environmental drivers of selection. We tested four landscape genomics methods on a simulated clinal population to determine their effectiveness at identifying a locus under varying selection strengths along an environmental gradient. We found all methods produced very low type I error rates across all selection strengths, but elevated type II error rates under "weak" selection. We then applied these methods to an AFLP genome scan of an alpine plant, Campanula barbata, and identified five highly supported candidate loci associated with precipitation variables. These loci * These authors contributed equally to this work.