Subtle Introgression Footprints at the End of the Speciation Continuum in a Clade of Heliconius Butterflies.

Quantifying gene flow between lineages at different stages of the speciation continuum is central to understanding speciation. Heliconius butterflies have undergone an adaptive radiation in wing color patterns driven partly by natural selection for local mimicry. Color patterns are also known to be used as assortative mating cues. Therefore, wing pattern divergence is considered to play a role in speciation. A corollary is that mimicry between closely related species may be associated with hybridization and interfere with reproductive isolation. Here, we take a multifaceted approach to explore speciation history, species boundaries, and traits involved in species differentiation between the two closely related species, Heliconius hecale and Heliconius ismenius. We focus on geographic regions where the two species mimic each other and contrast this with geographic regions where they do not mimic each other. To examine population history and patterns of gene flow, we tested and compared a four-population model accounting for linked selection. This model suggests that the two species have remained isolated for a large part of their history, yet with a small amount of gene exchange. Accordingly, signatures of genomic introgression were small except at a major wing pattern allele and chemosensing genes and stronger in the mimetic populations compared with nonmimetic populations. Behavioral assays confirm that visual confusion exists but that short-range cues determine strong sexual isolation. Tests for chemical differentiation between species identified major differences in putative pheromones which likely mediate mate choice and the maintenance of species differences.

Landscape context affects the sustainability of organic farming systems.

Organic agriculture promotes sustainability compared to conventional agriculture. However, the multifunctional sustainability benefits of organic farms might be mediated by landscape context. Assessing how landscape context affects sustainability may aid in targeting organic production to landscapes that promote high biodiversity, crop yields, and profitability. We addressed this using a meta-analysis spanning 60 crop types on six continents that assessed whether landscape context affected biodiversity, yield, and profitability of organic vs. conventional agroecosystems. We considered landscape metrics reflecting landscape composition (percent cropland), compositional heterogeneity (number and diversity of cover types), and configurational heterogeneity (spatial arrangement of cover types) across our study systems. Organic sites had greater biodiversity (34%) and profits (50%) than conventional sites, despite lower yields (18%). Biodiversity gains increased as average crop field size in the landscape increased, suggesting organic farms provide a "refuge" in intensive landscapes. In contrast, as crop field size increased, yield gaps between organic and conventional farms increased and profitability benefits of organic farming decreased. Profitability of organic systems, which we were only able to measure for studies conducted in the United States, varied across landscapes in conjunction with production costs and price premiums, suggesting socioeconomic factors mediated profitability. Our results show biodiversity benefits of organic farming respond differently to landscape context compared to yield and profitability benefits, suggesting these sustainability metrics are decoupled. More broadly, our results show that the ecological, but not the economic, sustainability benefits of organic agriculture are most pronounced in more intensive agricultural landscapes.

Balancing Bt Toxin Avoidance and Nutrient Intake by Helicoverpa zea (Lepidoptera: Noctuidae) Larvae.

To evaluate how the Cry1Ac Bacillus thuringiensis (Bt) toxin and the ratio of dietary protein to carbohydrate (P:C) independently and jointly affect Helicoverpa zea (Boddie) larval feeding performance and behavior, we conducted no-choice and binary choice experiments using chemically defined diets. We tested two related strains of this polyphagous pest: a strain selected for resistance to Cry1Ac in the laboratory (GA-R), and its field-derived parent strain (GA). In no-choice tests, feeding performance in GA and GA-R was superior on a non-Cry1Ac 80P:20C diet compared to a non-Cry1Ac 35P:65C diet or an 80P:20C diet containing Cry1Ac. Based on consumption and position measurements in binary choice tests, larvae preferred the non-Cry1Ac 80P:20C diet over the other two diets mentioned above. However, the association between preference and performance was weaker when comparing the Cry1Ac 80P:20C diet versus a Cry1Ac 35P:65C diet, and when comparing the Cry1Ac 80P:20C diet versus the non-Cry1Ac 35P:65C diet. In all choice situations, consumption preference occurred mainly by the percentage of larvae that fed almost entirely from one or the other diet, rather than from variation in the extent of diet mixing by individuals. Resistance to Cry1Ac affected the balance between toxin avoidance and nutrient intake: larvae from the more resistant GA-R strain consumed significantly more of the Cry1Ac 80P:20C diet when paired with non-Cry1Ac 35P:65C diet, while GA larvae consumed more of the latter, though not significantly. The results show that dietary P:C ratio, Cry1Ac, and resistance to Cry1Ac affected feeding behavior and performance of H. zea larvae.

Pear psylla and natural enemy thresholds for successful integrated pest management in pears.

Pear psylla, Cacopsylla pyricola (Förster), is the most economically challenging pest of commercial pears in Washington and Oregon, the top producers of pears in the United States. The objective of this study was to quantify economic injury levels and thresholds for pear psylla. We used the relationship between pear psylla adult and nymph densities, and fruit downgraded due to psylla honeydew marking to identify injury levels. We calculated economic injury levels using the cost of downgraded fruit and average management costs (spray materials and labor). Using economic injury levels, we determined economic thresholds for pear psylla, which include predicted pest population growth, natural enemy predation, and anticipated delays between when pest populations are measured and when managers apply interventions. Economic thresholds generated by this study were 0.1-0.3 second-generation nymphs per leaf and 0.2-0.8 third-generation nymphs per leaf depending on predicted price and yield for insecticide applications at 1,300 pear psylla degree days in the second generation and 2,600 pear psylla degree days in the third generation. Natural enemy inaction thresholds identified by this study were 6 Deraeocoris brevis or 3 Campylomma verbasci immatures per 30 trays or 2 earwigs per trap for third-generation optional insecticide applications.

Turning a Pest into a Natural Enemy: Removing Earwigs from Stone Fruit and Releasing Them in Pome Fruit Enhances Pest Control.

The European earwig Forficula auricularia (L.) (Dermaptera: Forficulidae) is an omnivorous insect that is considered a minor pest of stone fruit and a key predator of pests in pome fruit orchards. In many pome fruit orchards, earwigs are absent or in low abundance due to broad-spectrum spray programs and the slow recolonization rate of earwigs. Orchards in transition to organic or "selective" conventional programs often struggle to achieve effective levels of biological control, and thus, may benefit from inoculating earwigs to expedite their re-establishment. In a two-year study, we evaluated the potential for mass trapping earwigs from stone fruit using rolled cardboard traps to reduce fruit damage and provide earwigs for augmentation in pome fruit. We also tested whether a single mass release or five releases (on alternating weeks) of the same total number of earwigs in apples and pears reduced pests relative to plots where no releases occurred. Mass trapping did not decrease earwig abundance or substantially reduce fruit damage in stone fruit orchards. However, trapping was an efficient method for providing earwigs for augmentation. Earwig abundances were only increased in orchards where earwigs were previously low or absent; however, multiple orchards with varying prior levels of earwigs exhibited reductions in key pests (woolly apple aphid and pear psylla). For some other pests evaluated, plots with mass releases of earwigs had a slight trend in overall lower pest density when compared with control plots. A strategy for moving earwigs out of stone fruit orchards and into pome fruit orchards could be an effective method for augmenting orchard predator populations.

Seasonal activity of Trechnites insidiosus (Hymenoptera: Encyrtidae) and its host Cacopsylla pyricola (Hemiptera: Psyllidae) in pear.

Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae) is the most expensive and challenging insect pest of commercial pear trees in the Pacific Northwest. Integrated pest management (IPM) programs are working toward relying more heavily on natural enemies to reduce insecticide use. Trechnites insidiosus (Crawford) (Hymenoptera: Encyrtidae) is the main parasitoid of C. pyricola, but little is known about its biology in the region. Developing sampling tools is important for the deployment of IPM programs, including monitoring of natural enemies. In this study, we examined 2 conventional monitoring methods: beat trays and yellow sticky cards, in addition to screened sticky cards and 3D-printed cylinder traps. Additionally, we tested an overwintering trap for the collection of parasitized C. pyricola. The trapping methods were tested in orchards in Oregon and Washington. Unscreened cards caught the most T. insidiosus and C. pyricola, followed by screened cards, cylinder traps, and then beat trays. Beat trays sometimes failed to catch any T. insidiosus, even when it was found in abundance via other methods. Screened cards and cylinder traps reduced bycatch and increased ease of identifying T. insidiosus. Specimens from the cylinder traps were also more suitable for use in molecular analysis. The overwintering traps were effective at capturing parasitized C. pyricola, but were highly variable year to year. The ideal trapping method will vary based on research needs (e.g., DNA preservation, reducing bycatch, catching higher numbers), but both screened sticky cards and cylinder traps were viable methods for monitoring T. insidiosus and its host.

Integrated Pest Management Programs for Pear Psylla, Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae), Using Kaolin Clay and Reflective Plastic Mulch.

Pear psylla, Cacopsylla pyricola (Förster) (Hemiptera: Psyllidae), is the most economically important pest of pears grown in Washington State. Standard conventional management programs involve season-long broad-spectrum insecticide sprays. Although the industry uses some tools that are not disruptive to biological control, such as kaolin clay and selective insecticides, they are additions to broad-spectrum insecticides instead of replacements. Conventional sprays suppress pear psylla through the spring and early summer; however, disruption of biological control leads to pear psylla outbreaks near harvest. In 2018 and 2019, we tested two season-long programs that used only selective approaches. The programs began with either kaolin clay or reflective plastic mulch and were followed by identical spray programs using only selective insecticides. Programs were compared with an industry standard conventional program that used numerous broad-spectrum insecticides throughout the season, and a check program with no insecticides for pear psylla. Experiments were conducted using replicated 40-tree plots in a research orchard near Wenatchee, WA with high pear psylla pressure. In both years, selective programs had similar pear psylla densities to the industry standard program and all had lower pear psylla densities and fruit injury than the check. Both selective programs had lower fruit injury than the industry standard in the first year, and similar injury to the industry standard in the second year. Our results suggest kaolin clay and reflective mulch can effectively suppress pear psylla populations and injury in the early season and support season-long selective management programs without the use of broad-spectrum insecticides.

Effects of restricting movement between root and canopy populations of woolly apple aphid.

Movement of insect pests between spatially subdivided populations can allow them to recolonize areas where local extinction has occurred, increasing pest persistence. Populations of woolly apple aphid (Eriosoma lanigerum [Hausmann]; Hemiptera: Aphididae), a worldwide pest of apple (Malus domestica [Borkhausen]), occur both below- and aboveground. These spatially subdivided subpopulations encounter different abiotic conditions, natural enemies, and control tactics. Restricting movement between them might be an effective management tactic to decrease woolly apple aphid persistence and abundance. We examined this possibility in the field, using sticky barriers to restrict upward woolly apple aphid movement to tree canopies, and in the greenhouse, using mulches and sand amendments to restrict downward movement to roots. In the field, blocking aphid movement up tree trunks did not decrease the number of colonies in tree canopies. Instead, sticky-banded apple trees had higher aphid colony counts late in the study. Earwigs, which are woolly apple aphid predators, were excluded from tree canopies by sticky bands. In the greenhouse, fewer root galls (indicative of aphid feeding) occurred on trees in sandy potting media and on those with mulch (wood chips or paper slurry). Our results suggest that upward movement is less important than other factors that affect aboveground aerial woolly apple aphid population dynamics. In addition, apple orchards planted in sandier soils or with mulches may be partially protected from woolly apple aphid root feeding.