Adaptation without Specialization Early in a Host Shift.

AbstractStudents of speciation debate the role of performance trade-offs across different environments early in speciation. We tested for early performance trade-offs with a host shift experiment using a member of the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae). In this clade of plant-feeding insects, different species live on different host plants and exhibit strong behavioral and physiological host specialization. After five generations, the experimental host shifts resulted either in no adaptation or in adaptation without specialization. The latter result was more likely in sympatry; in allopatry, populations on novel host plants were more likely to become extinct. We conclude that in the early stages of speciation, adaptation to novel host plants does not necessarily bring about performance trade-offs on ancestral environments. Adaptation may be facilitated rather than hindered by gene flow, which prevents extinction. Additional causes of specialization and assortative mating may be required if colonization of novel environments is to result in speciation.

Eliminating explanations for Maladera formosae (Coleoptera: Scarabaeidae) preponderance in sandy soil.

Most field corn in the United States receives a neonicotinoid seed treatment for the management of early-season, soil-dwelling insect pests. Grubs of Maladera formosae (Brenske) (Coleoptera: Scarabaeidae) have been reported feeding on young field corn with both low and high rates of clothianidin seed treatments in Indiana, Michigan, and Ohio. Anecdotally, these infestations are restricted to sandy soils in the region. The purpose of this study was to (1) evaluate whether grub populations in corn are restricted to sandy soils, (2) assess whether soil type influences M. formosae survival, and (3) determine whether soil type affects clothianidin uptake by the plant, possibly explaining the observed differences in M. formosae abundance by soil type. We observed nearly 10-times more grubs in sand (>80% sand content) than loam (<80% sand content) soil within a single corn field. Grub survival to adult was not influenced by soil type. We then compared the concentrations of clothianidin seed treatment in the roots and shoots of corn seedlings grown in either sand or loam soil over time. Similar amounts of the active ingredient were found in the roots and shoots of corn grown in both soil types. Within 2 week, the clothianidin concentrations in both soil types had significantly declined in roots and shoots and were no different from the no-insecticide control. These findings suggest that factors other than insecticide exposure contribute to the higher abundance of M. formosae larvae in sand relative to loam soils, even within the same field.

Spatial distribution of Aphis glycines (Hemiptera: Aphididae): a summary of the suction trap network.

The soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), is an economically important pest of soybean, Glycine max (L.) Merrill, in the United States. Phenological information of A. glycines is limited; specifically, little is known about factors guiding migrating aphids and potential impacts of long distance flights on local population dynamics. Increasing our understanding of A. glycines population dynamics may improve predictions of A. glycines outbreaks and improve management efforts. In 2005 a suction trap network was established in seven Midwest states to monitor the occurrence of alates. By 2006, this network expanded to 10 states and consisted of 42 traps. The goal of the STN was to monitor movement of A. glycines from their overwintering host Rhamnus spp. to soybean in spring, movement among soybean fields during summer, and emigration from soybean to Rhamnus in fall. The objective of this study was to infer movement patterns of A. glycines on a regional scale based on trap captures, and determine the suitability of certain statistical methods for future analyses. Overall, alates were not commonly collected in suction traps until June. The most alates were collected during a 3-wk period in the summer (late July to mid-August), followed by the fall, with a peak capture period during the last 2 wk of September. Alate captures were positively correlated with latitude, a pattern consistent with the distribution of Rhamnus in the United States, suggesting that more southern regions are infested by immigrants from the north.

Aphid resistance is the future for soybean production, and has been since 2004: efforts towards a wider use of host plant resistance in soybean.

The soybean aphid (Aphis glycines) is an important pest of soybeans in the Midwestern US. The first aphid resistance genes were identified in the early 21st century and resistant varieties have been commercially available for 10 years, but have been very underutilized. Major seed companies have avoided commercializing aphid resistant soybean varieties for conventional farmers (i.e., not organic), in part because of the discovery of virulent biotypes in North America. The emergence of soybean aphid populations resistant to insecticides creates a greater incentive for the use of host plant resistance. New research on aphid genetics and markers, plant gene expression and in-plant refuges, suggest important avenues for insect resistance management (IRM) which may encourage more widescale commercialization of this valuable pest management tool.

Evaluation of Slug Refuge Traps in a Soybean Reduced-Tillage Cover Crop System.

As more farmers adopt no- or reduced-tillage and/or cover crop land management practices, slugs have become more frequent pests of field crops, including soybean. Monitoring slugs visually is difficult because they are nocturnal, so several trapping methods have been developed, though comparisons of trap types are rare. The objective of this study was to compare trapping efficiency of two types of slug refuge traps in reduced-tillage soybeans following cover crop termination. We tested a traditional shingle trap and a modified shingle trap with a water-filled pitfall trap beneath it. Traps were deployed in 24 pairs in 2018 and 2019 in experimental soybean plots. We counted slug captures weekly over a 5-week time period each year. In 2018, we counted the total number of slugs under each trap type. In 2019, counts were categorized into specific trap components (shingle vs. in/on/under the pitfall). Temperature was also recorded in 2019. The modified shingle traps captured significantly more slugs than the traditional shingle traps, mainly due to the pitfall component. This trend was most pronounced as slug density decreased, suggesting that the modified shingle trap is a more sensitive sampling tool which may be particularly valuable when used for research purposes.

Geographic Distribution of Bacillus thuringiensis Cry1F Toxin Resistance in Western Bean Cutworm (Lepidoptera: Noctuidae) Populations in the United States.

The western bean cutworm (WBC), Striacosta albicosta (Lepidoptera: Noctuidae), can be a severe pest of transgenic corn in the western Plains and Great Lakes regions of North America, including on hybrids expressing the Bacillus thuringiensis (Bt) Cry1F toxin. The level and geographic distribution of Cry1F resistance are not completely known. Neonate S. albicosta from 10 locations between Nebraska and New York state were subjected to dose-response trypsin-activated native Cry1F toxin overlay bioassays. In 2017, the mean estimated lethal concentration causing 50% larval mortality (LC50) ranged from 15.1 to 18.4 µg Cry1F cm-2, and were not significantly different among locations. In 2018, LC50 estimates at Scottsbluff, NE (22.0 µg Cry1F cm-2) and Watertown, NY (21.7 µg Cry1F cm-2) were significantly higher when compared to locations in Michigan (15.8 µg Cry1F cm-2). Significantly lower 14-day larval weight among survivors was correlated with higher Cry1F dose. Results from this study indicate that S. albicosta survivorship on purified Bt Cry1F toxin shows a relatively even distribution across the native and range expansion areas where seasonal field infestations typically occur.

Resistance of Soybean Plant Introductions to Three Colonies of Soybean Aphid (Hemiptera: Aphididae) Biotype 4.

Soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae), infestations of soybean, Glycine max (L.) Merr. (Fabales: Fabaceae), and the associated yield loss have led to a large dependence on insecticidal management in soybean throughout the Midwestern United States. However, several populations of pyrethroid-resistant soybean aphids have recently been found in Iowa, Minnesota, North Dakota and South Dakota, which highlights the importance of alternative management approaches. One such alternative method is host-plant resistance, which uses naturally occurring plant defenses in crop cultivars to reduce the potential for yield loss from a pest population. Current soybean aphid-resistant cultivars do not protect against all soybean aphids due to the presence of virulent biotypes. In particular, soybean aphid biotype 4 is virulent to Rag1 and Rag2 resistance genes both individually and in combination. However, we hypothesized that resistance to biotype 4 may exist in previously identified, but uncharacterized resistant soybean plant introductions (PIs). To test this, we evaluated 51 previously identified but uncharacterized soybean aphid-resistant PIs for their resistance to colonies of soybean aphid biotype 4 collected in separate site-years (Lomira, WI 2013; Volga, SD 2015, 2016). Free-choice tests identified 14 PIs with putative resistance to 'Lomira13', two to 'Volga15', and eight to 'Volga16' soybean aphid colonies. Follow-up, no-choice tests corroborated two to three resistant PIs per colony, and PI 437696, which was resistant to each of the three colonies and could aid in breeding efforts and an integrated approach to soybean aphid management.

Infestation ratings database for soybean aphid on early-maturity wild soybean lines.

Soybean aphid (Aphis glycines Matsumura; SA) is a major invasive pest of soybean [Glycine max (L.) Merr.] in northern production regions of North America. Although insecticides are currently the main method for controlling this pest, SA-resistant cultivars are being developed to sustainably manage SA in the future. The viability of SA-resistant cultivars may depend on identifying a diverse set of resistance genes from screening various germplasm sources, including wild soybean (Glycine soja Siebold and Zucc.), the progenitor of cultivated soybean. Data consisted of infestation ratings generated for a total of 337 distinct plant introduction lines of wild soybean that were exposed to avirulent SA biotype 1 for 14 d in 25 separate tests. Individual plants of the test lines were given a common rating by two researchers, based on a rating scale that progressed from 1=0 to 50, 2=51 to 100, 3=101 to 150, 4=151 to 200, 5=201 to 250, and 6 with >250 SA per test plant. Public dissemination of this dataset will allow for further analyses and evaluation of resistance among the test lines.

Biodiversity as both a cause and consequence of resource availability: a study of reciprocal causality in a predator-prey system.

1. One of the oldest questions in ecology is how species diversity in any given trophic level is related to the availability of essential resources that limit biomass (e.g. water, nutrients, light or prey). Researchers have tried to understand this relationship by focusing either on how diversity is influenced by the availability of resources, or alternatively, how resource abundance is influenced by species diversity. These contrasting perspectives have led to a seeming paradox '... is species diversity the cause or the consequence of resources that limit community biomass?' 2. Here we present results of an experiment that show it is possible for species diversity and resource density to exhibit reciprocal causal relationships in the same ecological system. Using a guild of ladybeetle predators and their aphid prey, we manipulated the number of predator species in field enclosures to examine how predator diversity impacts prey population size. At the same time, we manipulated the abundance of aphid prey in discrete habitat patches within each enclosure to determine how smaller-scale spatial variation in resource abundance affects the number of co-occurring predator species. 3. We found that the number of ladybeetle species added to enclosures had a significant impact on aphid population dynamics because interference competition among the predators reduced per capita rates of predation and, in turn, the overall efficiency of the predator guild. At the same time, spatial variation in aphid abundance among smaller habitat patches generated variation in the observed richness of ladybeetles because more species occurred in patches where predators aggregated in response to high aphid density. 4. The results of our experiment demonstrate that it is possible for species diversity to simultaneously be a cause and a consequence of resource density in the same ecological system, and they shed light on how this might occur for groups of mobile consumers that exhibit rapid responses to spatial and temporal variation in their prey.

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs.

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.