Diversity Of Herbivorous Insects Research Articles

Impact of vereda dryness on the insect herbivore diversity in adjacent cerrado areas in Brazilian Protected Areas

Unravelling the factors that influence plant-insect interactions remains a fundamental concern in terrestrial ecology. In the present study, we evaluated the impact of varying degrees of drying on cerrado areas adjacent to veredas on the species richness, abundance and composition of the insect herbivore fauna. Additionally, we examined differences in these impacts between various guilds (chewing and sucker insects) and developmental stages (adult and immature insects). The research has been performed in study areas located in three Protected Areas in Brazil. In total, we recorded 106 species of herbivorous insects across five orders (Coleoptera, Hemiptera, Hymenoptera, Lepidoptera, and Orthoptera). Hemiptera and Coleoptera stood out as the orders with the highest species diversity and individual numbers. Among the families, Cicadellidae and Curculionidae exhibited the highest richness in species and abundance of individuals. We found that richness of herbivorous insects was higher in wet zones (20.2 ± 6.5) of the veredas compared to dry zones (12.3 ± 7.5), as well as the abundance of herbivorous insects was higher in wet zones (49.4 ± 25.5) than in the dry zones (15.8 ± 7.9). Whithin insect guilds, the abundance of chewing insects was higher in wet zones (34.0 ± 13.6) than in dry zones (16.3 ± 14.8), but this difference was not observed for the diversity of sucking insects. Regarding developmental stage, immature insects exhibited higher species richness in wet zones (10.4 ± 5.0) compared to dry zones (5.3 ± 2.1), while adult insect diversity did not show differences between the studied zones. Furthermore, the species composition of herbivorous insects, various trophic guilds (chewers and suckers) and developmental stages (adults and immature insect) did not significantly differ among the zones studied within the veredas. According to our findings, herbivorous insects exhibit a preference for wet environments, which provide better conditions and higher-quality food resources for development and reproduction.

Effects of nitrogen inputs and mowing on the abundance and species richness of herbivorous insects in a meadow steppe.

To reveal the effects of nitrogen (N) inputs and mowing on the abundance and richness of insect community in meadow steppe, we investigated the abundance and diversity of herbivorous insects under four treatments in Inner Mongolia meadow steppe in August 2022, including control, N addition, mowing, and combined N addition and mowing. At a long-term control experimental platform, we collected insects using the vacuum sampling method. The results showed that N addition significantly increased the abundance of herbivorous insects, and mowing significantly decreased the abundance of herbivorous insects. Nitrogen addition significantly increased insect abundance in unmown condition but not in the mown condition. The responses of insect abundance at the community level to N addition were mainly driven by the dominant groups, Cicadellidae and Lygaeidae, which was affected by the abundance of Gramineae species. In contrast, their responses to mowing were the opposite. Both N addition and mowing did not affect the diversity of herbivorous insects. Our results indicated that the responses of herbivorous insect abundance to N inputs and mowing were directly regulated by food resources. With increasing food resource availability, the abundance but not the diversity of herbivorous insects increased, with stronger responses of dominant groups than subordinate ones.

Comparing the Above and Below-Ground Chemical Defences of Three Rumex Species Between Their Native and Introduced Provenances.

Compared to their native range, non-native plants often experience reduced levels of herbivory in the introduced range. This may result in reduced pressure to produce chemical defences that act against herbivores. We measured the most abundant secondary metabolites found in Rumex spp., namely oxalates, phenols and tannins. To test this hypothesis, we compared native (UK) and introduced (NZ) provenances of three different Rumex species (R. obtusifolius, R. crispus and R. conglomeratus, Polygonaceae) to assess whether any significant differences existed in their levels of chemical defences in either leaves and roots. All three species have previously been shown to support a lower diversity of insect herbivores and experience less herbivory in the introduced range. We further examined leaf herbivory on plants from both provenances when grown together in a common garden experiment in New Zealand to test whether any differences in damage might be consistent with variation in the quantity of chemical defences. We found that two Rumex species (R. obtusifolius and R. crispus) showed no evidence for a reduction in chemical defences, while a third (R. conglomeratus) showed only limited evidence. The common garden experiment revealed that the leaves analysed had low levels of herbivory (~ 0.5%) with no differences in damage between provenances for any of the three study species. Roots tended to have a higher concentration of tannins than shoots, but again showed no difference between the provenances. As such, the findings of this study provide no evidence for lower plant investments in chemical defences, suggesting that other factors explain the success of Rumex spp. in New Zealand.

Can the enemy release hypothesis explain the success of Rumex (Polygonaceae) species in an introduced range?

The enemy release hypothesis states that introduced plants have a competitive advantage due to their release from co-evolved natural enemies (i.e., herbivores and pathogens), which allows them to spread rapidly in new environments. This hypothesis has received mixed support to date, but previous studies have rarely examined the herbivore community, plant damage, and performance simultaneously and largely ignored below-ground herbivores. We tested for enemy release by conducting large scale field surveys of insect diversity and abundance in both the native (United Kingdom) and introduced (New Zealand) ranges of three dock (Rumex, Polygonaceae) species: R. conglomeratus Murray (clustered dock), R. crispus L. (curly dock) and R. obtusifolius L. (broad-leaved dock). We captured both above- and below-ground insect herbivores, measured herbivore damage, and plant biomass as an indicator for performance. In the introduced range, Rumex plants had a lower diversity of insect herbivores, all insect specialists present in the native range were absent and plants had lower levels of herbivore damage on both roots and leaves. Despite this, only R. crispus had greater fresh weight in the introduced range compared to the native range. This suggests that enemy release, particularly from below-ground herbivores, could be a driver for the success of R. crispus plants in New Zealand, but not for R. conglomeratus and R. obtusifolius.

Weak effects of birds, bats, and ants on their arthropod prey on pioneering tropical forest gap vegetation.

The relative roles of plants competing for resources versus top-down control of vegetation by herbivores, in turn impacted by predators, during early stages of tropical forest succession remain poorly understood. Here we examine the impact of insectivorous birds, bats, and ants exclusion on arthropods communities on replicated 5 × 5 m of pioneering early successional vegetation plots in lowland tropical forest gaps in Papua New Guinea. In plots from which focal taxa of predators were excluded we observed increased biomass of herbivorous and predatory arthropods, and increased density, and decreased diversity of herbivorous insects. However, changes in the biomass of plants, herbivores, and arthropod predators were positively correlated or uncorrelated between these three trophic levels and also between individual arthropod orders. Arthropod abundance and biomass correlated strongly with the plant biomass irrespective of the arthropods' trophic position, a signal of bottom-up control. Patterns in herbivore specialization confirm lack of a strong top-down control and were largely unaffected by the exclusion of insectivorous birds, bats, and ants. No changes of plant-herbivore interaction networks were detected except for decrease in modularity of the exclosure plots. Our results suggest weak top-down control of herbivores, limited compensation between arthropod and vertebrate predators, and limited intra-guild predation by birds, bats, and ants. Possible explanations are strong bottom-up control, a low activity of the higher order predators, especially birds, possibly also bats, in gaps, and continuous influx of herbivores from surrounding mature forest matrix.

Evolution of Olfactory Receptors Tuned to Mustard Oils in Herbivorous Drosophilidae.

The diversity of herbivorous insects is attributed to their propensity to specialize on toxic plants. In an evolutionary twist, toxins betray the identity of their bearers when herbivores coopt them as cues for host-plant finding, but the evolutionary mechanisms underlying this phenomenon are poorly understood. We focused on Scaptomyza flava, an herbivorous drosophilid specialized on isothiocyanate (ITC)-producing (Brassicales) plants, and identified Or67b paralogs that were triplicated as mustard-specific herbivory evolved. Using in vivo heterologous systems for the expression of olfactory receptors, we found that S. flava Or67bs, but not the homologs from microbe-feeding relatives, responded selectively to ITCs, each paralog detecting different ITC subsets. Consistent with this, S. flava was attracted to ITCs, as was Drosophila melanogaster expressing S. flava Or67b3 in the homologous Or67b olfactory circuit. ITCs were likely coopted as olfactory attractants through gene duplication and functional specialization (neofunctionalization and subfunctionalization) in S. flava, a recently derived herbivore.

Why species richness of plants and herbivorous insects do or do not correlate

AbstractUnraveling the determinants of herbivorous insect diversity has been a significant challenge in ecology. Despite the strong association between insect and plant species, previous studies conducted in natural systems have shown great variation in the strength of the correlation between their species richness. Such variation could be attributed to the proportion of generalist insect species (generality). However, both higher and lower generality may weaken the correlation because (a) generalist insect species are less dependent on the number of plant species, and (b) specialist insect species utilize only a proportion of the total plant species. To explore these contradictory effects, we studied plant and herbivorous insect communities in seminatural grasslands in Japan. Plant–insect interactions were evaluated in a unique way with a particular focus on the staying and herbivory behaviors of insects, which reflect their habitat use as well as host use. We found that lower generality of insect communities strengthened the correlation between species richness of plants and insects, but this was not the case when plant species that had no interaction with insect species were also considered. This was because lower generality increased the number of plant species that did not interact with insects. The results indicate that insect generality has contradictory effects on the plant–insect diversity relationship, which emphasizes the importance of distinguishing the effects to understand the variation in the relationship between plant and insect diversity in natural systems.

What We Don't Know About Diet-Breadth Evolution in Herbivorous Insects

Half a million species of herbivorous insects have been described. Most of them are diet specialists, using only a few plant species as hosts. Biologists suspect that their specificity is key to their diversity. But why do herbivorous insects tend to be diet specialists? In this review, we catalog a broad range of explanations. We review the evidence for each and suggest lines of research to obtain the evidence we lack. We then draw attention to a second major question, namely how changes in diet breadth affect the rest of a species’ biology. In particular, we know little about how changes in diet breadth feed back on genetic architecture, the population genetic environment, and other aspects of a species’ ecology. Knowing more about how generalists and specialists differ should go a long way toward sorting out potential explanations of specificity, and yield a deeper understanding of herbivorous insect diversity.

Contrasting Relationship of True Weevils (Curculionidae) and Woody Plants Diversity in Semi-arid Landscape Units.

In general, insect herbivore diversity is positively correlated to plant diversity since plant assemblages structure different microhabitats and provide food resources for such insects. Although poorly studied, insect herbivores in tropical dry forests are positively affected by tree species richness and the structural architecture of vegetation. In this study, we analyzed whether true weevil and woody plants present similar patterns of diversity across different landscape units in a Neotropical tropical dry forests located in Brazil, and if there is a correlation among their assemblages. We used Hill numbers according to species richness and the inverse of Simpson to compare the taxonomic diversity of true weevils and plants in nine landscape units located at the São Francisco river basin in the states of Sergipe and Alagoas, north-eastern Brazil. All trees and shrubs with diameter at breast height (1.3m) ≥ 5cm were sampled, and true weevils were collected using a modified Malaise trap. We used co-correspondence analysis to test if plants and true weevil species tend to co-occur. A total of 538 true weevils from 60 species was collected, and a total of 1419 plants belonging to 49 species was recorded. There is no general pattern of co-occurrence and diversity among true weevils and plants, indicating that the plant community is not structuring true weevil assemblage in the Caatinga. However, there was positive relationship between Sibinia sp4 and Fabaceae-Mimosoideae. These results suggest true weevil-plant relationships in the Caatinga differ from that of more mesic ecosystems. Therefore, we observed that the mechanisms that regulate herbivore-plant diversity relationships do not always follow a positive relationship, as observed in previous studies.

Insect herbivory in novel Quercus ilex L. forests: the role of landscape attributes, forest composition and host traits

Insect herbivory in novel Quercus ilex L. forests is a highly context- and scale-dependent process. We show that forest composition, tree height and fine-scale spatial location, as well as tree genetic relatedness and ontogeny, can all influence herbivore activity at local to landscape scale. Understanding the intrinsic and extrinsic drivers of herbivory in novel expanding forests is essential to envisage their role for biodiversity conservation. To analyse the effects of landscape attributes, forest composition, genetic relatedness, ontogeny and leaf traits on insect herbivory in novel Q. ilex forest stands. In 15 forest patches, we examined effects of patch size and connectivity, forest composition, tree height, specific leaf area (SLA) and nitrogen content on herbivory. In 3 forest patches, we assessed effects of tree genetic relatedness, ontogeny and spatial distribution. Herbivory was lower in pine-oak than in mixed-oak forests owing to the shorter tree height in the former with no effects of patch size or connectivity. Herbivory increased with SLA whereas nitrogen content had no effect. Within patches, herbivory differed among genetic clusters and was reduced in saplings growing near mature oaks and individuals near the forest edge. We illustrate the strong context and scale dependence of tree-herbivore interactions that renders predictions for dynamic systems such as novel oak forests extremely challenging. It implies, however, that the structural heterogeneity of such unmanaged forests allows their function as stepping stones for insect herbivore diversity in fragmented landscapes.

Common pheromone use among host-associated populations of the browntail moth, Euproctis chrysorrhoea, displaying different adult phenologies

The diversity of herbivorous insects may arise from colonization and subsequent specialization on different host plants. Such specialization requires changes in several insect traits, which may lead to host race formation if they reduce gene flow among populations that feed on different plants. Behavioural changes may play a relevant role in host race formation, for example if different races evolve distinct sexual communication signals or adult phenology. Previous research has revealed differences in larval phenology in different host-associated populations of the browntail moth, Euproctis chrysorrhoea (Lepidoptera: Erebidae). Here, sex pheromones among populations of this species are compared, and pheromone trapping data obtained is used in the field to build a phenological model that tests whether populations that feed on different plants differ in their adult flight period. The chemical and electrophysiological analyses revealed that two E. chrysorrhoea populations (on Prunus and on Arbutus unedo) use the same sex pheromone component for mate finding. Our trapping data, however, showed that males fly on average 25 days earlier in populations whose larvae feed on A. unedo compared to those whose larvae feed on Quercus species. Although the shifted phenology described here may underlie host-plant specialization in E. chrysorrhoea, and adults of this species are short-lived, the use of a common sexual pheromone and a large overlap in flight periods suggest that host race formation via allochronic isolation is unlikely in this moth. (Less)

Endemism in Wyoming plant and insect herbivore communities during the early Eocene hothouse

AbstractThe warm, equable, and ice-free early Eocene Epoch permits investigation of ecosystem function and macro-ecological patterns during a very different climate regime than exists today. It also provides insight into what the future may entail, as anthropogenic CO2release drives Earth toward a comparable hothouse condition. Studying plant–insect herbivore food webs during hothouse intervals is warranted, because these account for the majority of nonmicrobial terrestrial biodiversity. Here, we report new plant and insect herbivore damage census data from two floodplain sites in the Wind River Basin of central Wyoming, one in the Aycross Formation (50–48.25 Ma) at the basin edge (WRE) and the second in the Wind River Formation in the interior of the basin (WRI). The WRI site is in stratigraphic proximity to a volcanic ash that is newly dated to 52.416 ± 0.016/0.028/0.063 (2σ). We compare the Wind River Basin assemblages to published data from a 52.65 Ma floodplain flora in the neighboring Bighorn (BH) Basin and find that only 5.6% of plant taxa occur at all three sites and approximately 10% occur in both basins. The dissimilar floras support distinct suites of insect herbivores, as recorded by leaf damage. The relatively low-diversity BH flora has the highest diversity of insect damage, contrary to hypotheses that insect herbivore diversity tracks floral diversity. The distinctiveness of the WRE flora is likely due to its younger age and cooler reconstructed paleotemperature, but these factors are nearly identical for the WRI and BH floras. Site-specific microenvironmental factors that cannot be measured easily in deep time may account for these differences. Alternatively, the Owl Creek Mountains between the two basins may have provided a formidable barrier to the thermophilic organisms that inhabited the basin interiors, supporting Janzen's hypothesis that mountain passes appear higher in tropical environments.

CHANGES IN THE INSECT HERBIVORE FAUNA AFTER THE FIRST RAINS IN A TROPICAL DRY FOREST

Tropical dry forests are characterized by a pronounced dry season, when most trees shed their leaves, and a rainy season characterized by the production of new leaves. This study aimed to determine the effect of the first rains at the beginning of the rainy season on the insect herbivore fauna. We sampled 90 trees at the end of the dry season and 60 trees six days after the first rains using an entomological umbrella . Species richness and abundance of insect herbivores per tree was higher after the first rains. The results suggest a high synchrony between leaf production at the onset of the rainy season and the increase in insect herbivore diversity. Because young leaves are rich in nutrients and have a lower concentration of carbon-based defenses, many herbivore species in seasonal environments have their life cycles highly adjusted to the phenology of their host trees.

Mycorrhizal‐mediated plant–herbivore interactions in a high CO2 world

Abstract The symbiotic relationship between terrestrial plants and arbuscular mycorrhizal (AM) fungi is a key driver of plant nutritional and defence traits influencing insect herbivory. These tripartite interactions have been fundamental to shaping the evolution of land plants and the diversity of insect herbivores. Surprisingly, we have little understanding of how these interactions will function under elevated atmospheric CO2 concentrations (eCO2), despite the considerable implications for both natural and managed ecosystems. Although substantial research has revealed how eCO2 alters mycorrhizal–plant interactions, or plant–herbivore interactions, there is a stark scarcity of studies which investigate how eCO2 impacts mycorrhizal‐mediated plant–insect herbivore relationships. Here, we synthesise some of the main effects of eCO2 on the mycorrhizal symbiosis, the concomitant impacts on plant nutrient dynamics and secondary metabolism, and how eCO2‐driven changes in plant growth, biochemistry and communities impact insect herbivores. We point out that potential mechanistic drivers of AM fungal–plant–insect herbivore relationships under eCO2 can function antagonistically and are highly context‐dependent, which poses a particular challenge. Still, we hypothesise as to the potential outcomes for AM fungal–plant–herbivore dynamics under eCO2. We identify key research priorities to tackle the substantial gap in our understanding. If ecological theory is to effectively inform agricultural and natural management practices in the future, research needs to directly investigate how changes in global atmospheric CO2 concentrations impact the tripartite relationship between AM fungi, plants and insect herbivores. A plain language summary is available for this article.

Impact of urbanization on predator and parasitoid insects at multiple spatial scales.

Landscapes are becoming increasingly urbanized, causing loss and fragmentation of natural habitats, with potentially negative effects on biodiversity. Insects are among the organisms with the largest diversity in urbanized environments. Here, we sampled predator (Ampulicidae, Sphecidae and Crabronidae) and parasitoid (Tachinidae) flower-visiting insects in 36 sites in the city of Rome (Italy). Although the diversity of herbivorous insects in urban areas mostly depends on the availability of flowering plants and nesting sites, predators and parasitoids generally require a larger number of resources during their life cycle, and are expected to be particularly influenced by urbanization. As flower-visitors can easily move between habitat patches, the effect of urbanization was tested at multiple spatial scales (local, landscape and sub-regional). We found that urbanization influenced predator and parasitoid flower-visitors at all three spatial scales. At the local scale, streets and buildings negatively influenced evenness of predators and species richness and abundance of parasitoids probably acting as dispersal barrier. At the landscape scale, higher percentage of urban decreased predator abundance, while increasing their evenness, suggesting an increase in generalist and highly mobile species. Area and compactness (i.e. Contiguity index) of urban green interactively influenced predator communities, whereas evenness of parasitoids increased with increasing Contiguity index. At the sub-regional scale, species richness and abundance of predators increased with increasing distance from the city center. Compared to previous studies testing the effect of urbanization, we found little variation in species richness, abundance and evenness along our urbanization gradient. The current insect fauna has been probably selected for its tolerance to habitat loss and fragmentation, being the result of the intensive anthropogenic alteration occurred in the area in the last centuries. Conservation strategies aimed at predator and parasitoid flying insects have to take in account variables at multiple spatial-scales, as well as the complementarity of resources across the landscape.

Evolution of defences in large tropical plant genera: perspectives for exploring insect diversity in a tri-trophic context

Divergence and escalation in defences promote chemical diversity in plants, and consequently the diversity of insect herbivores. This diversification cascades to insect parasitoids through direct effects on host herbivore susceptibility, changes in herbivore community composition, or disparity in plant volatiles. Large tropical plant genera represent an ideal model for studying these trends due to the high diversity of sympatric species and their insects. Novel measures of chemical structural similarity should be used to analyse evolutionary trends in both direct and indirect defences. Host chemical data need to be combined with detailed herbivore and parasitoid data. This will help to identify truly active compounds. Furthermore, resolved genomic phylogenies for plants and insects should be included to assign directionality in the processes.

Unifying host-associated diversification processes using butterfly\u2013plant networks

Explaining the exceptional diversity of herbivorous insects is an old problem in evolutionary ecology. Here we focus on the two prominent hypothesised drivers of their diversification, radiations after major host switch or variability in host use due to continuous probing of new hosts. Unfortunately, current methods cannot distinguish between these hypotheses, causing controversy in the literature. Here we present an approach combining network and phylogenetic analyses, which directly quantifies support for these opposing hypotheses. After demonstrating that each hypothesis produces divergent network structures, we then investigate the contribution of each to diversification in two butterfly families: Pieridae and Nymphalidae. Overall, we find that variability in host use is essential for butterfly diversification, while radiations following colonisation of a new host are rare but can produce high diversity. Beyond providing an important reconciliation of alternative hypotheses for butterfly diversification, our approach has potential to test many other hypotheses in evolutionary biology.

Do Cultivars of Native Plants Support Insect Herbivores?

Native plants are becoming widely used in built landscapes to help mitigate the loss of biodiversity caused by urbanization. The primary advantage of native plant species over introduced ornamentals is their ability to support the development of the insects that fuel vertebrate food webs as well as specialist pollinators. The horticultural industry has introduced many cultivars of native plants to improve their aesthetic value and disease resistance, but there has been little work that measures the impact of these genetic changes on insect herbivores and pollinators. Here we measure how six desirable traits in native woody plant cultivars (leaf color, variegation, fall color, habit, disease resistance, and fruit size) compare with their wild types in terms of their ability to support insect herbivore development, abundance, and species richness. Using a common garden experiment, we quantified the abundance and diversity of insect herbivores using each species and its cultivars for growth and development over a 2-year period, as well as cumulative feed damage over the entire season. We also conducted feeding tests with evergreen bagworm (Thyridopteryx ephemeraeformis) to measure the preference of hatchling caterpillars for cultivars vs. straight species. We found that cultivars that had leaves altered from green to red, blue, or purple deterred insect feeding in all three experiments, a preference for variegated cultivars in one of our three experiments, but no consistent pattern of use among the species and cultivars chosen for other traits. These results suggest that the usefulness of native cultivars in restoring insect-driven food webs depends on the cultivar trait that has been selected.

Changing interactions among persistent species as the major driver of seasonal turnover in plant-caterpillar interactions.

β diversity of herbivorous insects in the tropics is usually very high, and there is often strong dissimilarity in herbivore species composition across different spatial scales and different abiotic gradients. Similarly, turnover is high for trophic interactions between herbivorous insects and their host plants. Two factors have been proposed to explain temporal or spatial differences in trophic interactions: changes in species composition and temporal changes in the behavior of shared species. The goal of this study was to evaluate determinants of high β diversity of trophic interactions between lepidopteran caterpillars and their host plants across dry and rainy seasons and their transitions. Over the course of a year, interaction diversity data were collected from 275 temporary plots in Cerrado vegetation, comprising 257 species of caterpillars, 137 species of host plants and 503 different trophic interactions. All these diversity parameters varied across seasons. Species assemblages of caterpillars and plants were different among the four seasons, and there was a high turnover of interactions between the seasons. The high temporal β diversity of trophic interactions was mostly due to interaction rewiring between co-occurring species, as opposed to changes in species composition over time.

Genetic drift precluded adaptation of an insect seed predator to a novel host plant in a long-term selection experiment.

Host specialization is considered a primary driver of the enormous diversity of herbivorous insects. Trade-offs in host use are hypothesized to promote this specialization, but they have mostly been studied in generalist herbivores. We conducted a multi-generation selection experiment to examine the adaptation of the specialist seed-feeding bug, Lygaeus equestris, to three novel host plants (Helianthus annuus, Verbascum thapsus and Centaurea phrygia) and to test whether trade-offs promote specialization. During the selection experiment, body size of L. equestris increased more on the novel host plant H. annuus compared to the primary host plant, Vincetoxicum hirundinaria, but this effect was not observed in other fitness related traits. In addition to selection, genetic drift caused variation among the experimental herbivore populations in their ability to exploit the host plants. Microsatellite data indicated that the level of within-population genetic variation decreased and population differentiation increased more in the selection line feeding on H. annuus compared to V. hirundinaria. We found a negative correlation between genetic differentiation and heterozygosity at the end of the experiment, suggesting that differentiation was significantly affected by genetic drift. We did not find fitness trade-offs between L. equestris feeding on the four hosts. Thus, trade-offs do not seem to promote specialization in L. equestris. Our results suggest that this insect herbivore is not likely to adapt to a novel host species in a time-scale of 20 generations despite sufficient genetic variation and that genetic drift disrupted the response to selection.