Morphometric criteria to differentiate Drosophila suzukii (Diptera: Drosophilidae) seasonal morphs.

Temperate insect species often enter diapause in preparation for overwintering. One such species is the invasive vinegar fly, Drosophila suzukii (Matsumura), which has seasonal polymorphisms, considered winter and summer morphs. To date, the morphs have been differentiated by color and size with winter morphs typically being darker and larger compared to summer morphs. 'Dark' and 'large' are subjective, however, and standardizing an identification process can ensure that the morph of interest is being accurately characterized. The goal of our research was to investigate a quantitative method to distinguish between D. suzukii morphs based on body and wing size. We reared winter and summer morph D. suzukii in the laboratory using standard procedures, and measured wing length, wing width, and hind tibia length. Additionally, we collected field D. suzukii to document the seasonal phenology of the morphs in Minnesota based on our model's cutoff criteria. A classification and regression tree analysis were used to determine which metrics would be best for predicting field-caught D. suzukii morphs. Using laboratory-reared flies as our known morphs for the training data in the classification model we developed classification trees based on wing length and the ratio of wing length to hind tibia length. The frequency of winter and summer morphs present in the field varied based on which classification tree was used. Nevertheless, we suggest ratio of wing length to hind tibia length as the most robust criteria for differentiating D. suzukii morphs because the ratio accounts for the size variability between laboratory-reared and field-caught flies and the error rate of misclassification is reduced to 0.01 for males. The results from this work can aid in future D. suzukii research by allowing scientists to objectively differentiate the morphs, and thereby improve our understanding of the biology and phenology of seasonal morph dynamics.

Proximate Drivers of Migration and Dispersal in Wing-Monomorphic Insects.

Gains in our knowledge of dispersal and migration in insects have been largely limited to either wing-dimorphic species or current genetic model systems. Species belonging to these categories, however, represent only a tiny fraction of insect biodiversity, potentially making generalization problematic. In this perspective, I present three topics in which current and future research may lead to greater knowledge of these processes in wing-monomorphic insects with limited existing molecular tools. First, threshold genetic models are reviewed as testable hypotheses for the heritability of migratory traits, using the sweet potato whitefly (Bemisia tabaci) as a case study of a behaviorally-polymorphic migratory species lacking morphological or physiological differentiation. In addition, both adaptive and non-adaptive explanations for the empirically variable relationship between egg production and flight in wing-monomorphic insects are discussed. Finally, with respect to the largest order of insects (Hymenoptera), the role of sex determination mechanisms for haplodiploidy as a driver for natal dispersal (for inbreeding avoidance) versus philopatry (such as in local mate competition) is discussed.

Dispersal strategies in terrestrial insects.

Terrestrial insects frequently disperse and/or migrate, either through their own self-directed actions or via other vehicles. Here, the following recent advances in the study of insect dispersal are highlighted: (1) components of classic hypotheses (marginal value theorem and inbreeding avoidance via sex-specific dispersal) have found varying degrees of recent support; (2) modern genetic tools have uncovered several candidate dispersal genes; (3) dispersal syndromes vary in their genetic and/or physiological constraints; and (4) common laboratory techniques may not accurately reflect dispersal in the field. A common theme is the tendency for breakthroughs to be concentrated in species with extremely well-defined dispersal phenotypes (e.g., long-distance migrants, wing polymorphic insects), suggesting the need for increased focus on species exhibiting less self-directed modes of dispersal.

Multistate Comparison of Attractants for Monitoring Drosophila suzukii (Diptera: Drosophilidae) in Blueberries and Caneberries.

Drosophila suzukii Matsumara, also referred to as the spotted wing drosophila, has recently expanded its global range with significant consequences for its primary host crops: blueberries, blackberries, raspberries, cherries, and strawberries. D. suzukii populations can increase quickly, and their infestation is difficult to predict and prevent. The development of effective tools to detect D. suzukii presence in new areas, to time the beginning of activity within a crop, to track seasonal activity patterns, and to gauge the effectiveness of management efforts has been a key research goal. We compared the efficiency, selectivity, and relationship to fruit infestation of a range of commonly used homemade baits and a synthetic formulated lure across a wide range of environments in 10 locations throughout the United States. Several homemade baits were more efficient than apple cider vinegar, a commonly used standard, and a commercially formulated lure was, in some configurations and environments, comparable with the most effective homemade attractant as well as potentially more selective. All alternative attractants also captured flies between 1 and 2 wk earlier than apple cider vinegar, and detected the presence of D. suzukii prior to the development of fruit infestation. Over half the Drosophila spp. flies captured in traps baited with any of the attractants were not D. suzukii, which may complicate their adoption by nonexpert users. The alternative D. suzukii attractants tested are improvement on apple cider vinegar and may be useful in the development of future synthetic lures.

Factors limiting the spread of the protective symbiont Hamiltonella defensa in Aphis craccivora Aphids.

Many insects are associated with heritable symbionts that mediate ecological interactions, including host protection against natural enemies. The cowpea aphid, Aphis craccivora, is a polyphagous pest that harbors Hamiltonella defensa, which defends against parasitic wasps. Despite this protective benefit, this symbiont occurs only at intermediate frequencies in field populations. To identify factors constraining H. defensa invasion in Ap. craccivora, we estimated symbiont transmission rates, performed fitness assays, and measured infection dynamics in population cages to evaluate effects of infection. Similar to results with the pea aphid, Acyrthosiphon pisum, we found no consistent costs to infection using component fitness assays, but we did identify clear costs to infection in population cages when no enemies were present. Maternal transmission rates of H. defensa in Ap. craccivora were high (ca. 99%) but not perfect. Transmission failures and infection costs likely limit the spread of protective H. defensa in Ap. craccivora. We also characterized several parameters of H. defensa infection potentially relevant to the protective phenotype. We confirmed the presence of H. defensa in aphid hemolymph, where it potentially interacts with endoparasites, and performed real-time quantitative PCR (qPCR) to estimate symbiont and phage abundance during aphid development. We also examined strain variation of H. defensa and its bacteriophage at multiple loci, and despite our lines being collected in different regions of North America, they were infected with a nearly identical strains of H. defensa and APSE4 phage. The limited strain diversity observed for these defensive elements may result in relatively static protection profile for this defensive symbiosis.

Worldwide populations of the aphid Aphis craccivora are infected with diverse facultative bacterial symbionts.

Facultative bacterial endosymbionts can play an important role in the evolutionary trajectory of their hosts. Aphids (Hemiptera: Aphididae) are infected with a wide variety of facultative endosymbionts that can confer ecologically relevant traits, which in turn may drive microevolutionary processes in a dynamic selective environment. However, relatively little is known about how symbiont diversity is structured in most aphid species. Here, we investigate facultative symbiont species richness and prevalence among world-wide populations of the cowpea aphid, Aphis craccivora Koch. We surveyed 44 populations of A. craccivora, and detected 11 strains of facultative symbiotic bacteria, representing six genera. There were two significant associations between facultative symbiont and aphid food plant: the symbiont Arsenophonus was found at high prevalence in A. craccivora populations collected from Robinia sp. (locust), whereas the symbiont Hamiltonella was almost exclusively found in A. craccivora populations from Medicago sativa (alfalfa). Aphids collected from these two food plants also had divergent mitochondrial haplotypes, potentially indicating the formation of specialized aphid lineages associated with food plant (host-associated differentiation). The role of facultative symbionts in this process remains to be determined. Overall, observed facultative symbiont prevalence in A. craccivora was lower than that of some other well-studied aphids (e.g., Aphis fabae and Acyrthosiphon pisum), possibly as a consequence of A. craccivora's almost purely parthenogenetic life history. Finally, most (70 %) of the surveyed populations were polymorphic for facultative symbiont infection, indicating that even when symbiont prevalence is relatively low, symbiont-associated phenotypic variation may allow population-level evolutionary responses to local selection.

Do trade-offs have explanatory power for the evolution of organismal interactions?

The concept of a trade-off has long played a prominent role in understanding the evolution of organismal interactions such as mutualism, parasitism, and competition. Given the complexity inherent to interactions between different evolutionary entities, ecological factors may especially limit the power of trade-off models to predict evolutionary change. Here, we use four case studies to examine the importance of ecological context for the study of trade-offs in organismal interactions: (1) resource-based mutualisms, (2) parasite transmission and virulence, (3) plant biological invasions, and (4) host range evolution in parasites and parasitoids. In the first two case studies, mechanistic trade-off models have long provided a strong theoretical framework but face the challenge of testing assumptions under ecologically realistic conditions. Work under the second two case studies often has a strong ecological grounding, but faces challenges in identifying or quantifying the underlying genetic mechanism of the trade-off. Attention is given to recent studies that have bridged the gap between evolutionary mechanism and ecological realism. Finally, we explore the distinction between ecological factors that mask the underlying evolutionary trade-offs, and factors that actually change the trade-off relationship between fitness-related traits important to organismal interactions.

Almost there: transmission routes of bacterial symbionts between trophic levels.

Many intracellular microbial symbionts of arthropods are strictly vertically transmitted and manipulate their host's reproduction in ways that enhance their own transmission. Rare horizontal transmission events are nonetheless necessary for symbiont spread to novel host lineages. Horizontal transmission has been mostly inferred from phylogenetic studies but the mechanisms of spread are still largely a mystery. Here, we investigated transmission of two distantly related bacterial symbionts--Rickettsia and Hamiltonella--from their host, the sweet potato whitefly, Bemisia tabaci, to three species of whitefly parasitoids: Eretmocerus emiratus, Eretmocerus eremicus and Encarsia pergandiella. We also examined the potential for vertical transmission of these whitefly symbionts between parasitoid generations. Using florescence in situ hybridization (FISH) and transmission electron microscopy we found that Rickettsia invades Eretmocerus larvae during development in a Rickettsia-infected host, persists in adults and in females, reaches the ovaries. However, Rickettsia does not appear to penetrate the oocytes, but instead is localized in the follicular epithelial cells only. Consequently, Rickettsia is not vertically transmitted in Eretmocerus wasps, a result supported by diagnostic polymerase chain reaction (PCR). In contrast, Rickettsia proved to be merely transient in the digestive tract of Encarsia and was excreted with the meconia before wasp pupation. Adults of all three parasitoid species frequently acquired Rickettsia via contact with infected whiteflies, most likely by feeding on the host hemolymph (host feeding), but the rate of infection declined sharply within a few days of wasps being removed from infected whiteflies. In contrast with Rickettsia, Hamiltonella did not establish in any of the parasitoids tested, and none of the parasitoids acquired Hamiltonella by host feeding. This study demonstrates potential routes and barriers to horizontal transmission of symbionts across trophic levels. The possible mechanisms that lead to the differences in transmission of species of symbionts among species of hosts are discussed.

Quantification and ultrastructure of oosorption in Eretmocerus eremicus (Hymenoptera: Aphelinidae).

The potential for and ultrastructure of oosorption were examined in Eretmocerus eremicus, a short-lived whitefly parasitoid that obligately produces anhydropic oocytes. In the absence of hosts, median egg load decreased by approximately 12% per day between 2 days and 8 days following eclosion. Parasitoid mating status had no significant effect on either egg load alone or the relationship between egg load and age. Yolk degradation in E. eremicus is autolytic, with the enzymes required for yolk sphere digestion apparently being derived from within the ooplasm. The exochorion appear to be digested by the follicular epithelium concurrent with the uniform degradation of the entire ooplasm. The potential adaptive benefits of this novel oosorption mechanism to E. eremicus females include a reduction in the total digestion time per oocyte and resorption of chorion remnants. Finally, to our knowledge, the results of this study provide the first unequivocal ultrastructural evidence of a purely autolytic oosorption mechanism in Insecta.