A global synthesis of the effects of diversified farming systems on arthropod diversity within fields and across agricultural landscapes.

Agricultural intensification is a leading cause of global biodiversity loss, which can reduce the provisioning of ecosystem services in managed ecosystems. Organic farming and plant diversification are farm management schemes that may mitigate potential ecological harm by increasing species richness and boosting related ecosystem services to agroecosystems. What remains unclear is the extent to which farm management schemes affect biodiversity components other than species richness, and whether impacts differ across spatial scales and landscape contexts. Using a global metadataset, we quantified the effects of organic farming and plant diversification on abundance, local diversity (communities within fields), and regional diversity (communities across fields) of arthropod pollinators, predators, herbivores, and detritivores. Both organic farming and higher in-field plant diversity enhanced arthropod abundance, particularly for rare taxa. This resulted in increased richness but decreased evenness. While these responses were stronger at local relative to regional scales, richness and abundance increased at both scales, and richness on farms embedded in complex relative to simple landscapes. Overall, both organic farming and in-field plant diversification exerted the strongest effects on pollinators and predators, suggesting these management schemes can facilitate ecosystem service providers without augmenting herbivore (pest) populations. Our results suggest that organic farming and plant diversification promote diverse arthropod metacommunities that may provide temporal and spatial stability of ecosystem service provisioning. Conserving diverse plant and arthropod communities in farming systems therefore requires sustainable practices that operate both within fields and across landscapes.

Warmer autumns and winters could reduce honey bee overwintering survival with potential risks for pollination services.

Honey bees and other pollinators are critical for food production and nutritional security but face multiple survival challenges. The effect of climate change on honey bee colony losses is only recently being explored. While correlations between higher winter temperatures and greater colony losses have been noted, the impacts of warmer autumn and winter temperatures on colony population dynamics and age structure as an underlying cause of reduced colony survival have not been examined. Focusing on the Pacific Northwest US, our objectives were to (a) quantify the effect of warmer autumns and winters on honey bee foraging activity, the age structure of the overwintering cluster, and spring colony losses, and (b) evaluate indoor cold storage as a management strategy to mitigate the negative impacts of climate change. We perform simulations using the VARROAPOP population dynamics model driven by future climate projections to address these objectives. Results indicate that expanding geographic areas will have warmer autumns and winters extending honey bee flight times. Our simulations support the hypothesis that late-season flight alters the overwintering colony age structure, skews the population towards older bees, and leads to greater risks of colony failure in the spring. Management intervention by moving colonies to cold storage facilities for overwintering has the potential to reduce honey bee colony losses. However, critical gaps remain in how to optimize winter management strategies to improve the survival of overwintering colonies in different locations and conditions. It is imperative that we bridge the gaps to sustain honey bees and the beekeeping industry and ensure food and nutritional security.

Age-based mating success in the codling moth, Cydia pomonella, and the obliquebanded leafroller, Choristoneura rosaceana.

In this study, the passage of spermatophores was examined between 1-day-old males mated in no-choice situations with females 0, 2, 4, or 6 days old and the converse for both the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), and the obliquebanded leafroller, Choristoneura rosaceana (Harris). For C. pomonella, female age had no effect on the passage of spermatophores from males, and only 6-day-old female C. rosaceana had reduced spermatophore number. The ages of moths at the time of mating had a greater effect on males, with C. pomonella males older than 2 days showing significant reductions in the ability to successfully pass a spermatophore to 1-day-old females. For C. rosaceana, 2- and 6-day-old males were significantly less likely to pass a spermatophore, but reduced transfer from 4-day-old males did not reach statistical significance. Wind-tunnel assays were used to evaluate the ability of 1- or 4-day-old males to fly upwind and successfully contact a young calling female. Four-day-old males were more likely to initiate flight upwind, but were less efficient at finding and contacting the females than younger males. This study suggests that evaluation of multiple components of the mating process are required to understand the effect of age at the time of mating on population dynamics of these moths.

Evaluating aggregation membership and copulatory success in the stink bug, Euschistus conspersus, using field and laboratory experiments.

The aggregation and mating behavior of the stink bug, Euschistus conspersus Uhler (Hemiptera: Pentatomidae) was investigated in a series of field and laboratory experiments. Marking of E. conspersus mating in aggregations in the field demonstrated that both sexes mate multiple times within aggregations on successive nights and with different partners, although ≈ 20% of the individuals of both sexes returned to aggregations but did not mate. Further analysis of mating patterns in caged aggregations revealed that heavy males and light females mated more frequently than their respective counterparts. Data are interpreted in terms of elucidating the function of benefits of multiple mating within aggregations for males and females.

Optimizing immunomarking systems and development of a new marking system based on wheat.

Immunomarking systems used to track large-scale movement patterns of insects are highly dependent on the efficiency of the enzyme linked imunosorbent assay (ELISA) reaction and logistical factors (e.g. concentration of marker applied, ability of the marker to wet the insect cuticle, and trapping methods). This paper examines ways to increase ELISA efficiency and mediate logistical factors, and provides information on a new immunomarking protein based on wheat gluten. The present studies on improving efficiency of the ELISA reactions showed that specially treated microplate surfaces were needed for soymilk and gluten assays, but not for egg albumin and casein assays. Sample dilution was investigated and was found to improve the signal/noise (S/N) ratio for the albumin and casein assays, but S/N ratios for the gluten and soymilk assays were less sensitive. However, for all assays, marked specimens were still detectable even with dilutions down to 6% of the original sample, which would allow more tests to be run on the same initial sample volume. For the logistical factors, these studies showed that marking of an insect by having it walk across a dried residue could be virtually eliminated for the casein and soymilk assays when the concentration applied was reduced to < 4%, but residues of 0.125% egg that had been aged in the field seven days still marked 37.5% of test insects placed on the residues. Also, the adjuvant Sylgard(®) 309 used at 80 ppm enhanced wetting of the insect cuticle and had little or no effect on the ELISA reaction, but the wetting agents R-11 and Silwet(®) L-77 were much more likely to negatively affect ELISA performance. Five different trapping adhesives were also evaluated and found to reduce ELISA efficiency 38-45% for the casein assay and 61-78% for the soymilk assay, while the albumin and gluten assays were unaffected. The information provided in this paper can be used to help correct for inherent differences in marking efficiency of the different proteins by manipulation of sample preparation, adjuvants, and concentrations applied.

Using phenology models to estimate insecticide effects on population dynamics: examples from codling moth and obliquebanded leafroller.

BACKGROUND: Phenology models based on degree-days were modified to estimate the effects of insecticide applications on the codling moth, Cydia pomonella (L.), and the obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae). Because the models inherently track stage structure of the population over the course of the season, stage specific mortality can be applied for various durations and intensities allowing the user to simulate different types of pesticides (ovicides, larvicides, organic or conventional materials). The models presented incorporate reproduction, effects of an application on subsequent generations, and the effects of each insecticide by itself as well as the combination of all the treatments. By comparing the treated populations to an untreated control, an estimate of how much the population size is reduced by different treatment programs over the entire season provides a clear assessment of the effectiveness of the treatment program. Simulations for codling moth also allow the insecticide effects to be simulated in conjunction with mating disruption (or not) and simulations for obliquebanded leafroller include the effect of insecticides on both instars 1-3 and 4-6. RESULTS: The simulations show there are distinct windows of opportunity for management programs, with rather large windows where an insecticide application has only a minor effect on population growth and also shows that intergenerational effects greatly affect optimal timings in subsequent generations. CONCLUSION: The models are especially useful in assessing current management strategies, testing potential improvements in timings, and evaluating cost savings and reducing non-target effects.

Recapture of codling moth (Lepidoptera: Tortricidae) males: influence of lure type and pheromone background.

Recapture of marked male codling moths, Cydia pomonella (L.) (Lepidoptera: Tortricidae), released four distances from traps was measured in experiments comparing either lure type or mating disruption. Experiment 1 assessed recapture by 0.1, 1, and 10 mg of codlemone lures. Experiments 2 and 3 assessed moth recapture in orchard plots with 0, 500, or 1,000 Isomate C Plus dispensers per ha. Moths were released 1, 3, 10, and 30 m downwind of the trap in experiments 1 and 2, and 3, 10, 30, and 45 m in experiment 3. Lure type did not affect recapture, however, significantly more moths were recaptured at 3 m compared with 10 or 30 m. Most moths recaptured < or = 10 m of the trap were recaptured by day 3, whereas most of the moths recaptured > or = 10 m were recaptured after day 3. Thus, 0.1-, 1-, and 10-mg lures, have an attractive range of between 10 and 30 m in orchards lacking mating disruption. Both mating disruption rates greatly reduced moth recapture, and moths recaptured under a 1,000 dispenser per ha rate were recaptured from < or = 10 m and within the first 2 d after release. Similar results were observed when release points were expanded to 45 m. Thus, results suggest that pheromone dispenser technologies and placement strategies that maximize disruption of males that arise within 10 m of a female are needed to markedly improve mating disruption.

Seasonal Population Dynamics of Potato Psyllid (Hemiptera: Triozidae) in the Columbia River Basin.

Understanding factors that affect the population dynamics of insect pest species is key for developing integrated pest management strategies in agroecosystems. Most insect pest populations are strongly regulated by abiotic factors such as temperature and precipitation, and assessing relationships between abiotic conditions and pest dynamics can aid decision-making. However, many pests are also managed with insecticides, which can confound relationships between abiotic factors and pest dynamics. Here we used data from a regional monitoring network in the Pacific Northwest United States to explore effects of abiotic factors on populations of an intensively managed potato pest, the potato psyllid (Bactericera cockerelli Sulc), which can vector Candidatus Liberibacter psyllaurus, a bacterial pathogen of potatoes. We assessed effects of temperature on psyllid populations, and show psyllid population growth followed predictable patterns within each year, but there was considerable variation across years in psyllid abundance. Examination of seasonal weather patterns suggested that in 2017, when psyllid populations were less abundant by several orders of magnitude than other years, a particularly long and cold period of winter weather may have harmed overwintering populations and limited population growth. The rate of degree-day accumulation over time, as well as total degree-day accumulation also affected trap catch abundance, likely by mediating the number of psyllid generations per season. Our findings indicate that growers can reliably infer the potential magnitude of risk from potato psyllids using monitoring data, date of first detection, seasonal weather patterns, and population size early in the growing season.

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.

Factors influencing mate choice in Euschistus conspersus Uhler (Heteroptera: Pentatomidae).

The patterns of mating and possible factors influencing mate choice in the consperse stink bug, Euschistus conspersus Uhler, were studied in a series of laboratory experiments. Males were found to transfer a significant percentage of their body mass during the initial mating. Mating was also found to reduce male longevity by 37.8% but had no significant effect on female longevity or fecundity. There was no evidence of male or female choice based on weight of potential mating partners. There was assortative mating based on experience, however, with males mating preferentially with virgin females and females preferring mated males when given the choice. The implications of these findings within the mating dynamics of this aggregating species are discussed.

Comparison of delayed female mating on reproductive biology of codling moth and obliquebanded leafroller.

Delay of mating was examined as a possible mechanism for population decreases associated with mating disruption for codling moth, Cydia pomonella L., and obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae). We examined the effect of delaying female mating 0, 2, 4, or 6 d while holding male age constant on life table parameters of both species. We found that increasing delays in mating were accompanied by two responses: (1) an increase in the percentage of sterile pairs and (2) a reduction in net reproductive rate and population growth unrelated to sterility. On a percentage basis, obliquebanded leafroller population growth was more strongly affected than codling moth. However, the net fertility rate of obliquebanded leafroller was nearly eight-fold higher than that of codling moth, so that obliquebanded leafroller females that experienced a 4-d delay in mating had nearly the same reproductive rate as codling moth females that experienced no delay. Leslie matrix simulations using life tables with field-based adult longevity estimates showed that codling moth females experiencing >2-d delay in mating resulted in decreases in population density or extinction within two generations. In contrast, obliquebanded leafroller females delayed <6 d showed rapid population growth that decreased as female age at mating increased; only the 6-d delay treatment resulted in decreased population levels. Our results indicate that obliquebanded leafroller females must on average experience a much longer delay in mating to significantly reduce population growth compared with codling moth females, suggesting that delay of mating likely plays a greater role in codling moth mating disruption than for obliquebanded leafroller.

Is biofix necessary for predicting codling moth (Lepidoptera: Tortricidae) emergence in Washington State apple orchards?

The heat-driven phenology model used for initiating codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), management in Washington state was examined to determine the need for using the capture of the first moth as a method of synchronizing the model and phenology of field populations (= biofix). We examined trap catch data taken at 1-2-d intervals from two research orchards; one data set encompassed a 28-yr period and the other data set a 4-yr period. We also examined consultant-collected data taken at 7-10-d intervals from 15 sites (N = 81), mostly between 2001 and 2005. At the two research sites, we found the mean biofix occurred at 96 degree-days (DD) (DD Celsius by using 10 degrees C lower threshold and 31.1 degrees C horizontal upper threshold) after 1 January (SD = 14.4; min. = 68, max = 122). After correcting for longer sampling intervals in the consultant data set, the biofix at the nonresearch sites occurred at 97 DD (N = 50, SD = 14.4; min. = 74, max = 120), nearly identical to that at the research sites. We also examined the performance of the codling moth model at predicting moth flight and egg hatch using a biofix and by just accumulating heat units from 1 January. The model performance was similar in both generations regardless of whether a biofix was used. The elimination of biofix simplifies management and eliminates mistakes associated with poor trap catch, particularly in low-pressure situations where mating disruption reduces trap efficiency.

Longevity of the adult codling moth, Cydia pomonella, and the obliquebanded leafroller, Choristoneura rosaceana, in Washington apple orchards.

The longevity of adult codling moth (Cydia pomonella (L.) Lepidoptera: Tortricidae) and obliquebanded leafroller (Choristoneura rosaceana (Harris) Lepidoptera: Tortricidae) held in shaded vials in the tree canopy was measured during the normal flight periods during 2004 and 2005. In both years all codling moths were dead by 130 degree-days (DD) (21 d) in the spring and 121 DD (8 d) in the summer. On a degree-day basis, data were similar across sex, generation, and year, and a common curve adequately predicted the survival distribution. For the obliquebanded leafroller, there were longevity differences between the sexes, but not between generations or years. Use of empirical quantile-quantile plots showed that the female obliquebanded leafroller lived an average of 32% longer than males. Maximum longevity observed in these studies for obliquebanded leafrollers was 117 DD (11 d) across both generations. The implications of these data for population biology studies and quarantine requirements are discussed.

Utilizing immunomarking techniques to track Halyomorpha halys (Hemiptera: Pentatomidae) movement and distribution within a peach orchard.

In this study we focus on the invasive brown marmorated stink bug, Halyomorpha halys (Stål) (Hemiptera: Pentatomidae), which has a strong dispersal capacity and has had a significant impact on several cropping systems, including peach (Prunus persica (L.)). Management of H. halys has relied on intensive insecticide use, and thus a better understanding of its dispersal behavior may assist in developing improved management strategies. In order to investigate H. halys movement and distribution patterns within a peach orchard we applied ecologically safe, food protein markers to the trees along the orchard border (chicken egg albumin in the form of liquid egg whites) and to the trees within the orchard interior (bovine casein in the form of cow's milk). We used enzyme-linked immunosorbent assays (ELISA) to assess whether collected H. halys were "marked" with either of the two protein markers, revealing where in the orchard the bugs had visited. From the density data we determined that H. halys is a perimeter-driven pest in peaches, with a significantly higher density of bugs collected along the orchard border. Interestingly, this trend is primarily driven by the distribution of male bugs. The protein marking data revealed that a small proportion of male H. halys move equally between the orchard border and interior, while a small proportion of females move predominately to the border after visiting the interior. The verification of a strong edge-effect, although potentially sex-specific, implies that H. halys displays a dispersal behavior that may also be exploited for management, which may help growers more efficiently and more effectively manage H. halys.

Evaluation of pheromone release from commercial mating disruption dispensers.

Pome fruit growers and crop consultants have expressed concerns about the seasonal release performance of commercial codling moth mating disruption dispenser products. Because of these concerns, we developed a laboratory flow-through volatile collection system (VCS) for measuring the volatile release of the codling moth sex pheromone, codlemone, from commercially available hand-applied dispensers. Under controlled air-flow and temperature conditions, the released vapor was trapped onto a polyurethane foam adsorbent followed by solvent extraction, solvent reduction, and GC/MS determination. Method recovery and breakthrough validations were performed to demonstrate system reliability before determining codlemone release from commercial dispensers field-aged over 140 days. The volatile collection was carried out in a consistent manner among five dispenser types most commonly used by growers, so that direct comparison of performance could be made. The comparison showed differences in the amount of pheromone released and in the patterns of release throughout the season between dispenser types. The variation in release performance demonstrates the need for routine evaluation of commercially marketed mating disruption dispensers. We believe that the simple and cost-effective volatile collection system can assist pheromone dispenser manufacturers in determining seasonal dispenser performance before new products are introduced into the commercial market and in rapidly verifying dispenser release when field-aged dispenser efficacy is in question.

Phenology of Lacanobia subjuncta (Lepidoptera: Noctuidae) in Washington and Oregon apple orchards.

The phenology of Lacanobia subjuncta (Grote & Robinson) (Lepidoptera: Noctuidae) was investigated in 30 apple orchards in central Washington state and northeastern Oregon from 1998 to 2001 (57 total orchard-yr). Adult captures in pheromone-baited traps were fit to a Weibull distribution to model emergence of the first and second generations. Initial capture of first generation adults was observed at 216.2 +/- 2.6 degree-days (DD) (mean +/- SEM) from 1 March by using a base temperature of 6.7 degrees C. The model predicted that flight was 5 and 95% complete by 240 and 700 degree-days (DD), respectively. Monitoring of oviposition and hatch was used to establish a protandry plus preoviposition degree-day requirement of 160.0 +/- 7.7 DD, as well as to provide data to describe the entire hatch period. Egg hatch was 5 and 95% complete by 395 and 630 DD, respectively. The start of the second flight was observed at 1217.1 +/- 8.3 DD by using an upper threshold for development of 32 degrees C and a horizontal cutoff. The model indicated that the second flight was 5 and 95% complete by 1220 and 1690 DD, respectively. Second generation hatch was 5 and 95% complete by 1440 and 1740 DD, respectively. A discussion of the potential uses of these detailed phenology data in optimizing management strategies is presented.

Instar-specific phenology of Pandemis pyrusana and Choristoneura rosaceana (Lepidoptera: Tortricidae) in Washington apple orchards.

Head capsule width was used to determine the instar specific phenology of the leafroller Pandemis pyrusana Kearfott and the obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), attacking apple in Washington state during 2001-2003. In total, 7012 P. pyrusana and 6122 obliquebanded leafroller larvae were measured from apple orchards from mid-March to mid-September. Degree-day accumulations from each site were paired with the head capsule data to determine the periods during which different instars were present in the field. The implications of this work for pest management and biological control of leafrollers is discussed.

A synthesis of the temperature dependent development rate for the obliquebanded leafroller, Choristoneura rosaceana.

The obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), is a serious pest of apples and other tree crops throughout North America. A review of temperature dependent development and models show that five different lower thresholds for development are published and used as the basis of heat-driven phenology models. We present a small lab data set of C. rosaceana development at four different temperatures and combine this with literature-based data into a single meta-analysis. Our analysis shows that the data from the different studies can be lumped together and the combined analysis suggests the lower and upper thresholds for development from egg to adult are approximately 10 degrees C and 30 degrees C, respectively.

Effect of Over-Tree Evaporative Cooling in Orchards on Microclimate and Accuracy of Insect Model Predictions.

Orchard design and management practices can alter microclimate and, thus, potentially affect insect development. If sufficiently large, these deviations in microclimate can compromise the accuracy of phenology model predictions used in integrated pest management (IPM) programs. Sunburn causes considerable damage in the Pacific Northwest, United States, apple-producing region. Common prevention strategies include the use of fruit surface protectants, evaporative cooling (EC), or both. This study focused on the effect of EC on ambient temperatures and model predictions for four insects (codling moth, Cydia pomonella L.; Lacanobia fruitworm, Lacanobia subjuncta Grote and Robinson; oblique-banded leafroller, Choristoneura rosaceana Harris; and Pandemis leafroller, Pandemis pyrusana Kearfott). Over-tree EC was applied in July and August when daily maximum temperatures were predicted to be ≥30°C between 1200-1700 hours (15/15 min on/off interval) in 2011 and between 1200-1800 hours (15/10 min on/off interval, or continuous on) in 2012. Control plots were sprayed once with kaolin clay in early July. During interval and continuous cooling, over-tree cooling reduced average afternoon temperatures compared with the kaolin treatment by 2.1-3.2°C. Compared with kaolin-treated controls, codling moth and Lacanobia fruitworm egg hatch in EC plots was predicted to occur up to 2 d and 1 d late, respectively. The presence of fourth-instar oblique-banded leafroller and Pandemis leafroller was predicted to occur up to 2 d and 1 d earlier in EC plots, respectively. These differences in model predictions were negligible, suggesting that no adjustments in pest management timing are needed when using EC in high-density apple orchards.

Fixed precision sampling plans for white apple leafhopper (Homoptera: Cicadellidae) on apple.

Constant precision sampling plans for the white apple leafhopper, Typhlocyba pomaria McAtee, were developed so that it could be used as an indicator species for system stability as new integrated pest management programs without broad-spectrum pesticides are developed. Taylor's power law was used to model the relationship between the mean and the variance, and Green's constant precision sequential sample equation was used to develop sampling plans. Bootstrap simulations of the sampling plans showed greater precision (D = 0.25) than the desired precision (Do = 0.3), particularly at low mean population densities. We found that by adjusting the Do value in Green's equation to 0.4, we were able to reduce the average sample number by 25% and provided an average D = 0.31. The sampling plan described allows T. pomaria to be used as reasonable indicator species of agroecosystem stability in Washington apple orchards.