The effectiveness of flower strips and hedgerows on pest control, pollination services and crop yield: a quantitative synthesis.

Floral plantings are promoted to foster ecological intensification of agriculture through provisioning of ecosystem services. However, a comprehensive assessment of the effectiveness of different floral plantings, their characteristics and consequences for crop yield is lacking. Here we quantified the impacts of flower strips and hedgerows on pest control (18 studies) and pollination services (17 studies) in adjacent crops in North America, Europe and New Zealand. Flower strips, but not hedgerows, enhanced pest control services in adjacent fields by 16% on average. However, effects on crop pollination and yield were more variable. Our synthesis identifies several important drivers of variability in effectiveness of plantings: pollination services declined exponentially with distance from plantings, and perennial and older flower strips with higher flowering plant diversity enhanced pollination more effectively. These findings provide promising pathways to optimise floral plantings to more effectively contribute to ecosystem service delivery and ecological intensification of agriculture in the future.

Bearing fruit: flower removal reveals the trade-offs associated with high reproductive effort for lowbush blueberry.

Past studies have shown that taxa from disparate groups often respond similarly to reduced reproductive effort. These common responses imply that high reproductive effort trades off with a consistent set of other life functions for most angiosperms, albeit modulated by their growth form and life history. However, many questions remain about reproductive trade-offs in plants, including just how many other life functions they involve, how diverse these functions may be, and how the severity of these trade-offs may vary through time. To address these questions in a long-lived, iteroparous shrub, we performed flower removal on plots of lowbush blueberry, Vaccinium angustifolium (Ericaceae), over 3 years. We found significant physiological differences between removal and control plots for ten diverse traits. Vegetative phenology was shifted earlier by about 20% in removal plots, and removal plots had about 15% more vegetative biomass by mid-season as well. Removal plots produced about 10% more ripe fruit per reproductive node by harvest than control plots, and reproductive nodes in removal plots produced at least one fruit by harvest about 6% more often. While fruit water content and titratable acidity were increased by removal, other fruit traits, such as sugar content and fresh mass, were not. The strength of the removal effect varied significantly by year for seven traits; for many, such as vegetative mass/stem and ripe fruit production/node, the effect was stronger in years with more stressful abiotic conditions. Our results demonstrate that there are tangible but variable costs to high reproductive effort for flowering plants.

Israeli acute paralysis virus: epidemiology, pathogenesis and implications for honey bee health.

Israeli acute paralysis virus (IAPV) is a widespread RNA virus of honey bees that has been linked with colony losses. Here we describe the transmission, prevalence, and genetic traits of this virus, along with host transcriptional responses to infections. Further, we present RNAi-based strategies for limiting an important mechanism used by IAPV to subvert host defenses. Our study shows that IAPV is established as a persistent infection in honey bee populations, likely enabled by both horizontal and vertical transmission pathways. The phenotypic differences in pathology among different strains of IAPV found globally may be due to high levels of standing genetic variation. Microarray profiles of host responses to IAPV infection revealed that mitochondrial function is the most significantly affected biological process, suggesting that viral infection causes significant disturbance in energy-related host processes. The expression of genes involved in immune pathways in adult bees indicates that IAPV infection triggers active immune responses. The evidence that silencing an IAPV-encoded putative suppressor of RNAi reduces IAPV replication suggests a functional assignment for a particular genomic region of IAPV and closely related viruses from the Family Dicistroviridae, and indicates a novel therapeutic strategy for limiting multiple honey bee viruses simultaneously and reducing colony losses due to viral diseases. We believe that the knowledge and insights gained from this study will provide a new platform for continuing studies of the IAPV-host interactions and have positive implications for disease management that will lead to mitigation of escalating honey bee colony losses worldwide.

Pesticide Contamination in Native North American Crops, Part I-Development of a Baseline and Comparison of Honey Bee Exposure to Residues in Lowbush Blueberry and Cranberry.

A pesticide exposure baseline for honey bees was compiled for two New England cropping systems, the native North American plant species consisting of lowbush blueberry (Vaccinium angustifolium Aiton) and cranberry (Vaccinium macrocarpon Aiton). More unique pesticide compounds were applied in blueberry than cranberry, but the numbers of pesticides discovered in trapped honey bee pollen were similar between the two crop systems. Not all pesticides found in pollen were the result of the applications reported by growers of either crop. When comparing residues, number of pesticides detected, total concentration, and risk quotient varied between the two crops. Also, blueberry was dominated by fungicides and miticides (varroacides) and cranberry was dominated by insecticides and herbicides. When comparing reported grower applications that were matched with detection in residues, the proportion of pesticide numbers, concentrations, and risk quotients varied by crop system and pesticide class. In most cases, pesticide residue concentrations were of low risk (low risk quotient) to honey bees in these crops. Estimation of decay rates of some of the most common pesticide residues under field conditions could aid growers in selection of less persistent compounds, together with safe application dates, prior to bringing in honey bees for pollination.

Pesticide Contamination in Native North American Crops, Part II-Comparison of Flower, Honey Bee Workers, and Native Bee Residues in Lowbush Blueberry.

In lowbush blueberry fields, we conducted residue analysis comparing flowers, trapped pollen (honey bee and Osmia spp.), and collected bees (honey bee workers, bumble bee queens, and non-Bombus spp. wild native bees). The study was conducted from 2012 to 2014. The number of pesticide residues, total concentrations, and risk to honey bees (Risk Quotient) on flowers were not significantly different from those determined for trapped honey bee pollen (except in one study year when residues detected in flower samples were significantly lower than residue numbers detected in trapped pollen). The compositions of residues were similar on flowers and trapped pollen. The number of residues detected in honey bee pollen was significantly greater than the number detected in Osmia spp. pollen, while the total concentration of residue was not different between the two types of pollen. The risk to honey bees was higher in trapped honey bee pollen than in trapped Osmia spp. pollen. The analysis of honey bee workers, native bumble bee queens, and native solitary bees showed that although more pesticide residues were detected on honey bee workers, there were no differences among the bee taxa in total residue concentrations or risk (as estimated in terms of risk to honey bees).

A deep multi-task learning approach to identifying mummy berry infection sites, the disease stage, and severity.

Introduction: Mummy berry is a serious disease that may result in up to 70 percent of yield loss for lowbush blueberries. Practical mummy berry disease detection, stage classification and severity estimation remain great challenges for computer vision-based approaches because images taken in lowbush blueberry fields are usually a mixture of different plant parts (leaves, bud, flowers and fruits) with a very complex background. Specifically, typical problems hindering this effort included data scarcity due to high manual labelling cost, tiny and low contrast disease features interfered and occluded by healthy plant parts, and over-complicated deep neural networks which made deployment of a predictive system difficult. Methods: Using real and raw blueberry field images, this research proposed a deep multi-task learning (MTL) approach to simultaneously accomplish three disease detection tasks: identification of infection sites, classification of disease stage, and severity estimation. By further incorporating novel superimposed attention mechanism modules and grouped convolutions to the deep neural network, enabled disease feature extraction from both channel and spatial perspectives, achieving better detection performance in open and complex environments, while having lower computational cost and faster convergence rate. Results: Experimental results demonstrated that our approach achieved higher detection efficiency compared with the state-of-the-art deep learning models in terms of detection accuracy, while having three main advantages: 1) field images mixed with various types of lowbush blueberry plant organs under a complex background can be used for disease detection; 2) parameter sharing among different tasks greatly reduced the size of training samples and saved 60% training time than when the three tasks (data preparation, model development and exploration) were trained separately; and 3) only one-sixth of the network parameter size (23.98M vs. 138.36M) and one-fifteenth of the computational cost (1.13G vs. 15.48G FLOPs) were used when compared with the most popular Convolutional Neural Network VGG16. Discussion: These features make our solution very promising for future mobile deployment such as a drone carried task unit for real-time field surveillance. As an automatic approach to fast disease diagnosis, it can be a useful technical tool to provide growers real time disease information that can prevent further disease transmission and more severe effects on yield due to fruit mummification.

The influence of insecticides and vegetation in structuring Formica mound ant communities (Hymenoptera: Formicidae) in Maine lowbush blueberry.

Assessing the influence of new, reduced-risk insecticides on natural enemies within agroecosystems is essential to developing integrated pest management strategies. Three species of mound-building Formica ants are abundant throughout Maine lowbush blueberry fields (Formica exsectoides Forel, F. glacialis Wheeler, and F. ulkei Emery). All three species have been described in the literature as predaceous, with research demonstrating that F. exsectoides preys on major pest insects of lowbush blueberry. The objectives of this study were to determine the impact of common-use and newly introduced insecticides on Formica sp. ant communities in lowbush blueberry fields. Laboratory assays indicated that the commonly applied insecticide phosmet is toxic to F. exsectoides, even after 8 d of field weathering (P < 0.05). Species comparisons indicated that susceptibility varied with exposure to residues in the field. However, some of the reduced-risk biorational insecticides, such as acetamiprid, had little effect on survival of all three species. Abundance of each species in the field varied with lowbush blueberry pesticide-use strategy and amount of nonblueberry vegetation. Both F. exsectoides and F. glacialis were most abundant in organic fields; however, overall F. glacialis was the most abundant in fields of all management types. Field surveys support laboratory results suggesting that phosmet is highly toxic to these species and influences their spatial pattern. Manipulation of the crop to conserve natural enemies in lowbush blueberry is difficult because the crop is not planted; therefore, we must look closely at the incorporation of low toxicity insecticides with natural enemies to efficiently control pest insects.

Monitoring of Spotted-Wing Drosophila (Diptera: Drosophilidae) Resistance Status Using a RAPID Method for Assessing Insecticide Sensitivity Across the United States.

Drosophila suzukii (Matsumura) has spread rapidly, challenging berry and cherry crop production due to its ability to lay eggs into ripening fruit. To prevent infestation by this pest, insecticides are applied during fruit ripening and harvest. We field-tested the Rapid Assessment Protocol for IDentification of resistance in D. suzukii (RAPID) on seventy-eight populations collected across eight U.S. states in 2017 and 2018. Exposure to LC50 rates of malathion, methomyl, spinetoram, spinosad, and zeta-cypermethrin led to average female fly mortality of 25.0% in 2017, and after adjusting concentrations the average was 39.9% in 2018. Using LC99 × 2 discriminating concentrations in 2017 and LC90 × 8 rates in 2018, average female mortalities were 93.3% and 98.5%, respectively, indicating high overall susceptibility. However, using these high concentrations we found 32.0% of assays with survival of some female flies in 2017 and 27.8% in 2018. The adjustment in discriminating dose from 2017 to 2018 also reduced the proportion of assays with <90% survival from 17.6 to 2.9%. Populations with low mortality when exposed to spinosad were identified using this assay, triggering more detailed follow-up bioassays that identified resistant populations collected in California coastal region berry crops. Widespread evaluations of this method and subsequent validation in California, Michigan, and Georgia in 2019-2021 show that it provides a quick and low-cost method to identify populations of D. suzukii that warrant more detailed testing. Our results also provide evidence that important insecticide classes remain effective in most U.S. regions of fruit production.

Honey Bee Health in Maine Wild Blueberry Production.

A two-year study was conducted in Maine wild blueberry fields (Vaccinium angustifolium Aiton) on the health of migratory honey bee colonies in 2014 and 2015. In each year, three or five colonies were monitored at each of nine wild blueberry field locations during bloom (mid-May until mid-June). Colony health was measured by assessing colony strength during wild blueberry bloom. Potential factors that might affect colony health were queen failure or supersedure; pesticide residues on trapped pollen, wax comb, and bee bread; and parasites and pathogens. We found that Varroa mite and pesticide residues on trapped pollen were significant predictors of colony health measured as the rate of change in the amount of sealed brood during bloom. These two factors explained 71% of the variance in colony health over the two years. Pesticide exposure was different in each year as were pathogen prevalence and incidence. We detected high prevalence and abundance of two recently discovered pathogens and one recently discovered parasite, the trypanosome Lotmaria passim Schwartz, the Sinai virus, and the phorid fly, Apocephalus borealis Brues.

Factors Influencing the Population Fluctuations of Euproctis chrysorrhoea (Lepidoptera: Erebidae) in Maine.

The browntail moth (Euproctis chrysorrhoea (L.)) is a forest pest that was accidentally introduced in the late 1800's and spread throughout New England in the early part of the 20th Century. At its peak range expansion in 1915 it encompassed an area of 150,000 km2 after which populations declined. By the 1960s, its distribution had receded to relic populations on outer Cape Cod, MA, and islands in Casco Bay, ME. In 1989 browntail moth resurged in Maine, with periodic, moderate outbreaks before a dramatic increase of the population occurred in 2016. We examined the pattern of annual defoliation by browntail moth since its resurgence in the 1990s as well as variation in populations throughout infested areas in Maine during three years of the recent outbreak, 2016-2018, relative to differences in weather, parasitism and habitat characteristics. Levels of defoliation over 24 yr were predicted by the preceding spring precipitation (-, negative effect) and the year's previous late summer and early fall temperatures (+, positive effect) when first to third instar larvae feed and then construct winter hibernacula. Late summer temperatures predicted the abundance of hibernacula across outbreak areas (+). Early spring temperatures (+) and early and late spring precipitation (-) predicted early summer larval and pupal nest abundance. Warmer fall temperatures result in more mature populations coming out of winter hibernacula in the spring, whereas spring precipitation drives epizootic outbreaks of Entomophaga aulicae (Reichardt in Bail) Humber (Entomophthorales: Entomophthoraceae). with parasitoids playing a lesser role. Climate trends indicate continued increases in fall temperatures since browntail moth resurgence.

Maine's Bumble Bees (Hymenoptera: Apidae)-Part 2: Comparisons of a Common (Bombus ternarius) and a Rare (Bombus terricola) Species.

As part of a quantitative survey of Maine's bumble bee fauna (Butler et al. 2021), we compared and contrasted genetic diversity, parasite and pathogen burdens, and pesticide exposure of the relatively common Bombus ternarius Say, 1937 and the spatially rare Bombus terricola Kirby, 1837. We recorded 11 Bombus species at 40 survey sites across three Maine ecoregions, and B. ternarius was the most common species, while B. terricola was spatially rare. Nonmetric multidimensional scaling indicated that B. terricola was associated with higher elevation sites in Maine, while B. ternarius was more broadly distributed in the state. Pollinator networks constructed for each bee indicated B. ternarius foraged on more plant species than B. terricola, but that there was considerable overlap (73%) in plant species visited. Genetic diversity was greater in the spatially restricted B. terricola, whereas the widely distributed B. ternarius was characterized by greater genetic differentiation among regions. Bombus terricola had higher molecular marker levels of the microsporidian fungi Nosema spp. and the trypanosome Crithidia spp., and both species had high levels of Trypanosoma spp. exposure. No Western Honey Bee (Apis mellifera, Linnaeus, 1758) viruses were detected in either species. Pesticides were not detected in pollen samples collected from workers of either species, and B. ternarius worker tissue samples exhibited only trace levels of diflubenzuron.

Maine's Bumble Bee (Hymenoptera: Apidae) Assemblage-Part 1: Composition, Seasonal and Regional Distribution, and Resource Use.

Global declines of bumble bees place natural and agricultural ecosystems at risk. Given bumble bees importance to Maine's major agricultural crops, we conducted a statewide, quantitative survey of bumble bee species seasonal and ecoregional abundance, richness, diversity, and floral resource use. We recorded 11 Bombus species at 40 survey sites across Maine's three ecoregions, with Bombus ternarius Cresson, 1863 and Bombus impatiens Cresson, 1863 being the most common and Bombus citrinus Smith, 1854 the least commonly encountered. Bumble bee species richness did not differ as a function of ecoregion, but did decline over the season, while species diversity differed by ecoregion and also declined over the season. Multiple response permutation procedure (MRPP) indicated ecoregional differences in species composition of bumble bee assemblages and nonmetric multidimensional scaling produced a stable ordination suggesting assemblage differences were associated with survey site variables including forage plant cover, forage plant richness, elevation, development, and deciduous forest cover. Both MRPP and correspondence analysis also revealed differences in the floral resources utilized by bumble bee species in each ecoregion. Low connectance and nestedness levels indicated low stability pollinator networks in each ecoregion, suggesting Maine bumble bee assemblages may be at risk of decline in response to additional external perturbations.

Field and Laboratory Testing of Feeding Stimulants to Enhance Insecticide Efficacy Against Spotted-Wing Drosophila, Drosophila suzukii (Matsumura).

The invasive spotted-wing drosophila, Drosophila suzukii (Matsumura), is a key insect pest of berries globally, causing lost revenues and increased production costs associated with applications of insecticides. The insecticides utilized are commonly broad-spectrum pyrethroids, organophosphates, or carbamates in conventionally managed fields and spinosad in organically managed fields. Adoption of more selective insecticides has been limited due to their lower residual activity, and the requirement that some must be ingested to be effective. We investigated the use of feeding stimulants for D. suzukii as a method to improve longevity and efficacy in a range of insecticides. In laboratory bioassays, sugar increased the efficacy of all chemical classes tested; however, the inclusion of yeast only showed a benefit with malathion. Feeding stimulants had a limited effect in some cases under field conditions. Similarly, infestation in field plots and a semifield bioassay showed no significant decreases in infestation with the inclusion of feeding stimulants for the insecticides tested in these trials. We discuss the implications of these findings for managing D. suzukii in fruit crops to help ensure the harvest of marketable fruit.

Population genomics of Drosophila suzukii reveal longitudinal population structure and signals of migrations in and out of the continental United States.

Drosophila suzukii, or spotted-wing drosophila, is now an established pest in many parts of the world, causing significant damage to numerous fruit crop industries. Native to East Asia, D. suzukii infestations started in the United States a decade ago, occupying a wide range of climates. To better understand invasion ecology of this pest, knowledge of past migration events, population structure, and genetic diversity is needed. In this study, we sequenced whole genomes of 237 individual flies collected across the continental United States, as well as several sites in Europe, Brazil, and Asia, to identify and analyze hundreds of thousands of genetic markers. We observed strong population structure between Western and Eastern US populations, but no evidence of any population structure between different latitudes within the continental United States, suggesting that there are no broad-scale adaptations occurring in response to differences in winter climates. We detect admixture from Hawaii to the Western United States and from the Eastern United States to Europe, in agreement with previously identified introduction routes inferred from microsatellite analysis. We also detect potential signals of admixture from the Western United States back to Asia, which could have important implications for shipping and quarantine policies for exported agriculture. We anticipate this large genomic dataset will spur future research into the genomic adaptations underlying D. suzukii pest activity and development of novel control methods for this agricultural pest.

Field Perimeter Trapping to Manage Rhagoletis mendax (Diptera: Tephritidae) in Wild Blueberry.

Rhagoletis mendax Curran (the blueberry maggot fly) is a major pest of wild blueberry. It is a direct pest of the fruit. Females lay eggs in fruit resulting in infestations of larvae unacceptable to most consumers. Three field perimeter interception tactics were tested for control of R. mendax in wild blueberry, Vaccinium angustifolium Aiton (Ericales: Ericaceae), between 2000 and 2010. We investigated field perimeter deployment of baited, insecticide-treated, green spheres (2000-2005), baited, yellow Trécé PHEROCON AM traps (2005 and 2006), and baited, Hopper Finder, sticky barrier tape (2008-2010). Only the Hopper Finder tape provided significant reduction in R. mendax adults and fruit infestation over the 3-yr field study. However, the reduction in fruit infestation compared with control plots was only 48.2 ± 7.3%, a level of reduction in damage that would be unsuitable for many commercial wild blueberry growers as a stand-alone tactic, but could be an important reduction as part of a multiple tactic IPM strategy. In addition, we constructed an agent-based computer simulation model to assess optimal trap placement between three patterns: 1) a single row of traps along field perimeter; 2) a double row of traps along the field perimeter at half the density of the single row; and 3) a grid of traps spread throughout the field but with the largest distance between traps. We found that the single row deployment pattern of traps was the best for reducing immigration of R. mendax adults into simulated fields.

Dispersal From Overwintering Sites, Action Thresholds for Blueberry Maggot Fly (Diptera: Tephritidae), and Factors That Can Influence Variation in Predicted Fruit Infestation Levels in Maine Wild Blueberry: Part I.

Between 1996 and 2003 field studies were conducted in wild blueberry, Vaccinium angustifolium Aiton (Ericales: Ericaceae) to determine action thresholds for the blueberry maggot fly, Rhagoletis mendax Curran (Diptera: Tephritidae). Thresholds were based upon cumulative fly trap captures on baited Pherocon AM traps. The cumulative numbers of R. mendax flies captured over time was a significant predictor of infested fruit levels (maggots/liter blueberries). Fifty percent of the variance in fruit infestation is explained by fly captures. Based upon this relationship, the University of Maine Cooperative Extension has recommended action threshold of 10 cumulative fly captures/trap. However, this threshold is liberal in terms of risk as only 50% of the variance in fruit infestation is explained by fly captures. The dynamics of colonization rate and fly physiological status entering fruit-bearing fields might partially explain the variance in the action threshold. The majority of flies emerge from pruned fields or along forest scrub/shrub field edges, and it takes between 1 and 10 d (mean = 4.1 ± 0.9 d) for the populations to enter adjacent fruit-bearing fields. Flies dispersed from pruned fields at a decreasing rate the farther they dispersed. The rate of dispersal into fruit-bearing wild blueberry fields also depends upon the overwintering site quality. Sites with fruit exhibited both delayed colonization and lower colonization rates into fruit-bearing fields than adjacent pruned fields with no fruit. We also found that as the season progressed the proportion of flies capable of laying eggs varied greatly by location and year.

Temporal Dynamics of the Blueberry Maggot Fly (Diptera: Tephritidae) and its Primary Parasitoid, Biosteres melleus (Hymenoptera: Braconidae), in the Maine Wild Blueberry System.

Between 1998 and 2017, we conducted studies in wild blueberry, Vaccinium angustifolium Aiton (Ericales: Ericaceae), to elucidate the temporal dynamics of the blueberry maggot fly, Rhagoletis mendax Curran, and its parasitoid, Biosteres melleus (Gahan). A predictive model for the emergence of R. mendax was validated at two sites over 3 yr. A second predictive model for the major parasitoid, B. melleus, of R. mendax was constructed and suggests that the delay in emergence of the parasitoid relative to its host provides a period or 'biological window' of 9 d where insecticide sprays can be applied to manage R. mendax with a limited impact on the parasitoid. A 20-yr study on the parasitoid/host dynamics showed parasitism rates ranging from 0.5 to 28.2%. It appears that R. mendax populations in Maine wild blueberry are characterized by stable equilibrium dynamics, significantly affected by stochastic processes. There was a weak, but significant relationship between B. melleus density and R. mendax intrinsic rates of growth. Our data suggest that R. mendax population dynamics in wild blueberry is characterized by an unstable equilibrium tipping point of 7.9 maggots per liter of blueberries or an average of 10 flies per trap.

Evaluation of Four Plant Extract Repellents for Management of the European Red Ant Myrmica rubra (Hymenoptera: Formicidae).

Spearmint oil, peppermint oil, neem oil, and d-limonene were tested as nest site repellents against the colonization of the invasive European red ant, Myrmica rubra (L.) in both laboratory and field trials. In a laboratory assay, a 10% (v/v) solution of each extract repelled M. rubra colonies from nesting in plant pots filled with moist soil compared to water-treated controls, when applied as a dip to pots. Extracts also repelled colonies compared to a water control in a second laboratory experiment, where pots were dipped 15 d prior to the start of the experiment. In a 2008 field comparison, 20 plant pots, filled with moist potting soil, were dipped in either 10% (v/v) spearmint oil or peppermint oil, 70% (v/v) neem oil, or a water control and left in infested sites. Seven control pots were colonized over 3 mo, while none of the extract-treated pots were colonized. This field trial was repeated for a 15-wk duration in 2009 with the addition of a 10% (v/v) d-limonene solution as a treatment, and a change in neem oil concentration to 10% (v/v). Spearmint and peppermint oils repelled M. rubra colonies for the duration of the experiment. Neem oil and d-limonene repelled colonies for 3 and 4 wk, respectively. These extracts, especially mint oils, show potential as low-hazard repellents against M. rubra in greenhouse and nursery settings, and could reduce the number of new infestations incurred by the transport of plant stock.

Movement of Rhagoletis mendax (Diptera: Tephritidae) in Fruit-Bearing Wild Blueberry Fields. Part II.

During dispersal into fruit-bearing wild blueberry fields, blueberry maggot flies were highly active during all daylight hours as revealed by trap captures, although in one trial afternoon activity was greater than morning activity. Flies were not captured in traps at night, although observations in growth chambers showed that their activity at night, measured as displacement of position, was equal to daylight conditions. Flies were shown to fly at low altitude, just above the crop canopy, and screen fencing was shown to be effective at reducing colonization of plots, presumably due to their low height during flight. Over a 4-yr mark-capture study, colonization rate was shown to be low at 9.7 m/d, although a separate 2010 study showed higher rates at 14.1 and 28.0 m/d. Movement was shown to be nondirectional or random in the field, but a constrained random walk exhibiting direction into the field. Weed cover and high fruit density were associated with higher fly relative abundance, suggesting these field characteristics served as attractors slowing colonization rate into a field. Transect trap studies showed the temporal and spatial pattern of fly colonization into commercial wild blueberry fields, one of a slow wave that penetrates into the field interior as the season progresses. There is also an increase in fly abundance within-field edges and adjacent forest. The 'stacking' of flies along a field edge and slow movement rate into a field was shown through simulation to be a result of nondirectional short-distance dispersal of flies.