A survey of syrphid predators of Nasonovia ribisnigri in organic lettuce on the central coast of California

Organic lettuce, Lactuca sativa L., producers on California's Central Coast rely on endemic syrphid flies (Diptera: Syrphidae) to suppress populations of Nasonovia ribisnigri Mosley (Homoptera: Aphididae) and other aphids affecting lettuce. Growers are using various forms of habitat manipulation to enhance biological control. We surveyed syrphids collected from organic romaine in and around the Salinas Valley from March through September 2005 to gain a better understanding of the species responsible for aphid suppression and to examine possible implications for biocontrol. The primary species of syrphid fly reared were Toxomerus marginatus (Say) (39%), Platycheirus stegnus (Say) (27%), Sphaerophoria sulfuripes (Thomson) (13%), and Allograpta obliqua (Say) (10%). Syrphus opinator Osten Sacken (2%), Toxomerus occidentalis (Curran) (1.3%), and Eupeodes volucris Osten Sacken (1%) were less common. Sphaerophoria pyrrhina Bigot, Scaeva pyrastri (L.), Platycheirus obscurus Say, Allograpta exotica Wiedemann, and Eupeodes americanus Wiedemann each made up <1% of the syrphids reared. T. marginatus and Sphaerophoria sulfuripes were commonly collected from romaine plants with few or no detectable aphids. P. stegnus was observed to deposit in clusters of eggs, and was only reared in significant numbers from highly infested fields. Approximately 5% of syrphid larvae overall were parasitized by either Diplazon sp. (Hymenoptera: Ichneumonidae) or Pachyneuron sp. (Hymenoptera: Pteromalidae).     

Syrphid flies suppress lettuce aphids

Syrphid flies are abundant in lettuce fields, where their larvae are key predators of aphids. However, the presence of predators in the field does not always result in economically significant levels of prey suppression. Even when predators are numerous, their effects on prey population dynamics may be variable. Over a two year period we surveyed lettuce fields in coastal California, USA to test whether syrphid flies are capable of colonizing fields with aphids and suppressing aphid population growth. The survey showed that female syrphids oviposited more eggs at locations with more aphids, and that greater numbers of syrphid larvae resulted in lower rates of increase in the aphid populations. We also directly manipulated syrphid densities by adding syrphid eggs to uncaged lettuce plants, and these syrphid additions resulted in lower aphid population growth. This research shows that syrphid flies have the ability to suppress aphid populations in lettuce fields.     

Oxygenated phosphine fumigation for control of Nasonovia ribisnigri (Homoptera: Aphididae) on harvested lettuce

Low temperature regular phosphine fumigations under the normal oxygen level and oxygenated phosphine fumigations under superatmospheric oxygen levels were compared for efficacy against the aphid, Nasonovia ribisnigri (Mosley), and effects on postharvest quality of romaine and head lettuce. Low temperature regular phosphine fumigation was effective against the aphid. However, a 3 d treatment with high phosphine concentrations of > or = 2,000 ppm was needed for complete control of the aphid. Oxygen greatly increased phosphine toxicity and significantly reduced both treatment time and phosphine concentration for control of N. ribisnigri. At 1,000 ppm phosphine, 72 h regular fumigations at 6 degrees C did not achieve 100% mortality of the aphid. The 1,000 ppm phosphine fumigation under 60% O2 killed all aphids in 30 h. Both a 72 h regular fumigation with 2,200 ppm phosphine and a 48 h oxygenated fumigation with 1,000 ppm phosphine under 60% O2 were tested on romaine and head lettuce at 3 degrees C. Both treatments achieved complete control of N. ribisnigri. However, the 72 h regular fumigation resulted in significantly higher percentages of lettuce with injuries and significantly lower lettuce internal quality scores than the 48 h oxygenated phosphine fumigation. Although the oxygenated phosphine fumigation also caused injuries to some treated lettuce, lettuce quality remained very good and the treatment is not expected to have a significant impact on marketability of the lettuce. This study demonstrated that oxygenated phosphine fumigation was more effective and less phytotoxic for controlling N. ribisnigri on harvested lettuce than regular phosphine fumigation and is promising for practical use.     

Growth, development and consumption by four syrphid species associated with the lettuce aphid, Nasonovia ribisnigri, in California

The lettuce aphid, Nasonovia ribisnigri Mosley, was accidentally introduced into California from Europe during the late 1990s and soon became an economic pest of Romaine lettuce along California’s central coast region. Indigenous syrphid larvae attack the lettuce aphid and are believed to be effective in the management of this invasive pest, although there have been no studies on the capacity of the syrphid larvae to kill and consume lettuce aphids. We focused on four syrphid species commonly found in central coast lettuce fields: Allograpta obliqua (Say), Eupeodes fumipennis (Thomson), Sphaerophoria sulphuripes (Thomson), and Toxomerus marginatus (Say). Laboratory feeding experiments were conducted to estimate the development times of all juvenile stages, the daily growth rate of larvae, the number of third instar aphids killed, the aphid biomass killed, and the aphid biomass consumed as measures of predator performance. Results show that during larval development E. fumipennis killed the most third-instar aphids (507 aphids, 88 mg biomass killed) and reached the largest size, followed by A. obliqua (228 aphids, 39 mg killed), S. sulphuripes (194 aphids, 31 mg killed) and T. marginatus (132 aphids, 20 mg killed). Body size alone did not account for species differences in per-capita larval consumption rates. This information is discussed in relation to the predation potential of syrphids through the short cropping cycle of lettuce, and the choice of plant species to use for floral resource provisioning to enhance the activity of syrphids needed for effective management of lettuce aphids in California’s central coast fields.     

Distribution and population development of Nasonovia ribisnigri (Homoptera: Aphididae) in iceberg lettuce

A field study was conducted to determine the distribution and development of aphid Nasonovia ribisnigri (Mosley) (Homoptera: Aphididae) populations in iceberg lettuce, Lactuca sativa L. 'Salinas'. Lettuce plants were transplanted and caged individually in the field and inoculated with apterous N. ribisnigri at 0, 1, 2, 3, and 4 wk after transplanting in spring and fall 2002. Plants were harvested 15-50 d after inoculations; numbers of alates and apterous N. ribisnigri were counted or estimated on each leaf for each plant. Inoculations during all 5 wk of plant development resulted in successful colonization of lettuce heads. Results indicated that head formation did not reduce the risk of colonization by N. ribisnigri to iceberg lettuce; plants were susceptible to colonization by N. ribisnigri throughout their development. For later inoculations, N. ribisnigri populations were relatively smaller, and aphids were found mostly within the heads. For earlier inoculations, N. ribisnigri populations were larger, and within-plant distributions shifted toward frame leaves. The shift of population distributions toward frame leaves correlated significantly with increases in N. ribisnigri population density. For most inoculations, more aphids were present on wrapper leaves than on other leaves. The proportion of alates did not vary significantly with population density. Population development of N ribisnigri also correlated significantly with heat unit accumulation. Yellow sticky cards were used to monitor alates in each cage. Catches of N. ribisnigri alates on yellow sticky cards were significantly correlated with total numbers of alates as well as with total population sizes on individual lettuce plants.     

Effects of Aphis fabae Scop, and of its attendant ants and insect predators on yields of field beans (Vicia faba L.)

Predators (mainly coccinellid adults and larvae and syrphid larvae), although few, were important in decreasing numbers of Aphis fabae on a small plot of field beans during the early stages of infestation in a year favourable to the aphid. At the same time, ants (Lasius niger L.), attending aphids on other plants on the same plot, effectively protected the aphids from predators for about 2 weeks, enabling the attended aphids to multiply faster than the unattended. When all aphid populations started to decline, predators became more numerous and accelerated the decline on both sets of plants. Bean plants without aphids yielded fifty-six seeds per plant; those with aphids but free from ants gave seventeen; and those with ant-attended aphids, eight seeds per plant. The damage and loss of yield was caused by the large aphid populations that developed when the pods were maturing, and not by the fewer aphids present when the plants were in flower. It appears that small, temporary infestations during flowering might increase the yield of field beans.     

Feeding and egg distribution studies of Heringia calcarata (Diptera: Syrphidae), a specialized predator of woolly apple aphid (Homoptera: Eriosomatidae) in Virginia apple orchards

Predation by the aphidophagous syrphid fly Heringia calcarata (Loew) on woolly apple aphid, Eriosoma lanigerum (Hausmann), was studied in the laboratory and in Virginia apple orchards. Feeding studies compared the prey suitability of three temporally sympatric aphid pests of apple: spirea aphid, Aphis spiraecola Patch; rosy apple aphid, Dysaphis plantaginea (Passerini); and woolly apple aphid. Significantly more H. calcarata larvae survived and completed development on a pure diet of woolly apple aphid than on rosy apple aphid, and none survived on spirea aphid. Final larval weights were significantly greater, and the larval developmental period was significantly shorter on woolly apple aphid than on rosy apple aphid, but neither the duration of pupal development nor adult weight differed between diets. H. calcarata larvae consumed an average of 105 woolly apple aphids during their development. Na?ve, neonate larvae given access to all possible pair combinations of woolly apple aphid, rosy apple aphid, and spirea aphid consumed significantly more woolly apple aphids in all pairings that included woolly apple aphid. When given a choice of rosy apple aphid and spirea aphid, significantly more rosy apple aphids were consumed. Weekly counts of syrphid eggs found in woolly apple aphid, rosy apple aphid, and spirea aphid colonies collected from apple trees showed that two generalist hover fly predators, Eupeodes americanus (Wiedemann) and Syrphus rectus Osten Sacken, were present in colonies of all three aphid species and that E. americanus was the most abundant syrphid predator in A. spiraecola and D. plantaginea colonies. H. calcarata eggs were found only in woolly apple aphid colonies and were more abundant there than E. americanus and S. rectus. These data suggest that H. calcarata is a specialized predator of woolly apple aphid in the apple ecosystem in Virginia.     

Effect of hunger on the allocation of time among pea plants by the larvae of an aphidophagous hover fly,Eupeodes corollae [Dipt: Syrphidae]

We observed the movement of predatory larvae of the syrphid flyEupeodes corollae (F.) (formerlyMetasyrphus corollae) among small pea plants with and without aphids. Starved larvae spent longer time than well-fed larvae on similar plants and both groups of larvae stayed longer on plants with aphids than on plants without aphids. On plants with aphids, larvae which failed to capture prey left the plant sooner than those which captured aphids. The capture of at least one aphid on a plant increased the persistence of syrphid larvae. The average rate of energy gain was higher for well-fed larvae than for starved larvae because starved larvac stayed on plants even when their rate of return was lower. When larvae that had captured aphids left plants, their rate of energy gain, tended to be lower than at any time following capture of the 2^nd, aphid. The 1^st aphid was captured in less time than similar larvae spent on plants without aphids. Time between captures of aphids by well-fed larvae was less than the time such larvae spent on plants without aphids. Among starved larvae, the intercatch intervals were similar to the time on plants without aphids. We discuss the significance of these results relative to current predator foraging theory and the efficiency ofE. corollae as a biological control agent.      

Do functional traits improve prediction of predation rates for a disparate group of aphid predators?

Aphid predators are a systematically disparate group of arthropods united on the basis that they consume aphids as part of their diet. In Europe, this group includes Araneae, Opiliones, Heteroptera, chrysopids, Forficulina, syrphid larvae, carabids, staphylinids, cantharids and coccinellids. This functional group has no phylogenetic meaning but was created by ecologists as a way of understanding predation, particularly for conservation biological control. We investigated whether trait-based approaches could bring some cohesion and structure to this predator group. A taxonomic hierarchy-based null model was created from taxonomic distances in which a simple multiplicative relationship described the Linnaean hierarchies (species, genera, etc.) of fifty common aphid predators. Using the same fifty species, a functional groups model was developed using ten behavioural traits (e.g. polyphagy, dispersal, activity, etc.) to describe the way in which aphids were predated in the field. The interrelationships between species were then expressed as dissimilarities within each model and separately analysed using PROXSCAL, a multidimensional scaling (MDS) program. When ordinated using PROXSCAL and then statistically compared using Procrustes analysis, we found that only 17% of information was shared between the two configurations. Polyphagy across kingdoms (i.e. predatory behaviour across animal, plant and fungi kingdoms) and the ability to withstand starvation over days, weeks and months were particularly divisive within the functional groups model. Confirmatory MDS indicated poor prediction of aphid predation rates by the configurations derived from either model. The counterintuitive conclusion was that the inclusion of functional traits, pertinent to the way in which predators fed on aphids, did not lead to a large improvement in the prediction of predation rate when compared to the standard taxonomic approach.     

Effects of vacuum and controlled atmosphere treatments on insect mortality and lettuce quality

Laboratory studies were conducted to determine the effects of vacuum and controlled atmosphere on mortality of aphids, Nasonovia ribisnigri (Mosley) and Macrosiphum euphorbiae (Thomas), and leafminer, Liriomyza langei Frick, and on the visual quality of iceberg lettuce at three different temperatures. Vacuum at 50 mbar and controlled atmosphere with 6% CO2 were effective in controlling aphids and leafminer larvae. Complete control of N. ribisnigri and M. euphorbiae was achieved with vacuum treatments and 6% CO2 CA treatments at 5 degrees C in 4 d. Mortality was >96% when leafminer larvae were treated with vacuum and 6% CO2 CA treatments for 4 d. However, leafminer pupae were more tolerant to the treatments and highest mortality was close to 60% in 4 d with CO2 under vacuum. None of the treatments had negative effects on visual quality of iceberg lettuce. Results from this study are encouraging and warrant further and large-scale research.     

The effect of co-infestation by conspecific and heterospecific aphids on the feeding behaviour of <Emphasis Type="Italic">Nasonovia ribisnigri</Emphasis> on resistant and susceptible lettuce cultivars

Aphid saliva can suppress the blocking of sieve elements, a reaction that plants employ to inhibit aphid feeding, but aphid saliva can also elicit plant defence responses. Such plant responses might affect interactions between different aphid species and intraspecifically, e.g. among different biotypes. The objectives of our study were to investigate if feeding behaviour and performance of two biotypes of the lettuce aphid Nasonovia ribisnigri are affected by (1) feeding by the other biotype and (2) feeding by the green peach aphid Myzus persicae or the potato aphid Macrosiphum euphorbiae. Additionally the effect of feeding in a group was studied. All experiments were performed on both a resistant and an isogenic susceptible lettuce cultivar. Feeding or probing by conspecific or heterospecific aphids had different effects on Nasonovia ribisnigri biotypes. Aphids were only slightly affected by feeding or probing of the same biotype on both susceptible and resistant lettuce. N. ribisnigri virulent biotype Nr:1 suppressed the resistance against Nr:0 in the resistant cultivar. In contrast, defence was induced by Nr:1 against Nr:0 in susceptible lettuce. Co-infestation by M. euphorbiae and M. persicae had minor effects on Nr:0. Defence against Nr:1 was induced on both susceptible lettuce and resistant lettuce by Nr:0 and M. euphorbiae. Additionally, M. persicae induced defence in resistant lettuce against Nr:1. Effectors in the saliva of Nr:1 aphids are likely responsible for the defence suppression in lettuce. Identification of these effectors could lead to a better understanding of the mechanism of virulence in N. ribisnigri.     

Agronomic aspects of strip intercropping lettuce with alyssum for biological control of aphids

Organic lettuce growers in California typically use insectary strips of alyssum (Lobularia maritima (L.) Desv.) to attract hoverflies (Syrphidae) that provide biological control of aphids. A two year study with transplanted organic romaine lettuce in Salinas, California investigated agronomic aspects of lettuce monoculture and lettuce-alyssum strip intercropping on beds in replacement intercropping treatments where alyssum transplants replaced 2 to 8% of the lettuce transplants, and in additive intercropping treatments where alyssum transplants were added to the standard lettuce density without displacing lettuce transplants. Alyssum and lettuce dry matter (DM) were determined at lettuce maturity. Alyssum transplants produced less shoot DM in the additive than in the replacement intercropping treatments. The number of open inflorescences of alyssum increased with alyssum DM, and among treatments ranged from 2 to15 inflorescences per lettuce head. Compared with monoculture lettuce, lettuce heads on intercropped beds were slightly smaller and had lower nitrogen concentrations in the both additive treatments and in some replacement treatments. This research provides the first information on a novel additive intercropping approach to provide alyssum floral resources for biological control of lettuce aphids, and suggests that this approach may be a more land-efficient particularly for producing smaller lettuce heads for romaine hearts or for markets with less strict size requirements. Additional research is needed to determine if the increased competition between alyssum and lettuce in additive intercropping would reduce lettuce yields for wholesale markets with larger head size requirements. Practical aspects of implementing the various intercropping arrangements and alternatives are discussed.     

Responses of Nasonovia ribisnigri (Homoptera: Aphididae) to susceptible and resistant lettuce

Nymphs and alates of aphid Nasonovia ribisnigri (Mosley) (Homoptera: Aphididae) were tested on 10 lettuce cultivars with N. ribisnigri resistance gene Nr and 18 cultivars without the resistance gene in various bioassays. Bioassays used whole plants, leaf discs, or leaf cages to determine susceptibility of commercial lettuce cultivars to N. ribisnigri infestation and to evaluate screening methods for breeding lettuce resistance to N. ribisnigri. Resistant and susceptible plants were separated in 3 d when using whole plant bioassays. Long-term (> or =7 d) no-choice tests using leaf cages or whole plants resulted in no survival of N. ribisnigri on resistant plants, indicating great promise of the Nr gene for management of N. ribisnigri. Effective screening was achieved in both no-choice tests where resistant or susceptible intact plants were tested separately in groups or individually and in choice tests where susceptible and resistant plants were intermixed. Leaf discs bioassays were not suitable for resistance screening. All lettuce cultivars without the resistance gene were suitable hosts for N. ribisnigri, indicating the great importance of this pest to lettuce production and the urgency in developing resistant lettuce cultivars to manage N. ribisnigri.     

Larval interactions in aphidophagous predators: effectiveness of wax cover as defence shield of Scymnus larvae against predation from syrphids

Small-sized predators in the aphidophagous guild of Aphis gossypii Glover colonies on hibiscus trees in Japan exploit aphids at low prey abundance. Scymnus (Pullus) posticalis Sicard beetles were the first predatory species to attack aphids in the spring, and their larvae co-occurred with larvae of Eupeodes freguens (Matsumura) syrphids in aphid-infested leaves of hibiscus for 3 weeks in absence of large-sized coccinellid predators. Larval interaction between Scymnus and syrphid predators was examined in relation to effectiveness of wax cover of Scymnus against predation from syrphids. Waxless first instar larvae were not protected but wax-covered larvae of second, third and fourth instars were protected from predation by syrphid larvae. The protection was lower in the second instar which has a thin wax cover and significantly higher in the third and fourth instars having a thick wax cover. In addition, larvae from which the wax was removed were significantly more vulnerable to predation. Vulnerability of Scymnus larvae to predation from syrphids was directly related to the thickness of wax cover. Results suggest that the wax cover of Scymnus larvae act as an effective defence mechanism against predation from syrphid larvae.     

The potential of predators in natural control of aphids in wheat: Results of a ten-year field study in two German landscapes

In the present study, we investigated the natural control of aphids by predators in wheat fields in a low (L) and high-input cropping region (H) of Germany during a 10-year period. Data for the statistical analyses were obtained from weekly after the start of aphid emergence. The mean annual aphid indices, calculated as the sum of Sitobion avenae (Fabr.), Rhopalosiphum padi (L.), Metopolophium dirhodum (Walk.)(Homoptera: Aphididae), were 30.4 and 81.5 × 103 aphid days per m2, for L and H, respectively. Nine predator fractions were analysed: Coccinella septempunctata L., adults (1) and larvae (2), Propylea quatuordecimpunctata (L.) (Coleoptera: Coccinellidae) adults (3) and larvae (4), syrphid larvae (mostly Episyrphus balteatus [De Geer] (Diptera: Syrphidae)) (5), Chrysoperla carnea Steph. (Neuroptera: Chrysopidae) larvae (6), and adult carabids (7), staphylinids (8) and spiders (9). The two sites were comparable in terms of the mean size of the overall predator community, expressed in predator units (PU): 4.9 PU/m2 (L) vs. 5.4 PU/m2 (H). Most predator fractions responded numerically to increasing aphid densities. The numerical response was strongest in syrphid larvae, scarcely detectable in adult coccinellids, and virtually non-existent in epigeic arthropods. Multiple regression models revealed indirect relationships between the weekly overall predator community densities (PU/m2) and individual predator fractions (individuals/m2) and absolute rates of aphid density increase (individuals/m2) one or two weeks after baseline. A site-independent reduction of the aphid density increase to nil (y = 0) was observed at 3.9 to 4.2 PU/m2. Consequently, the 2.7 times higher aphid density at H cannot be attributed to the presence of fewer predators or lower effects of the overall predator community or of any individual predator fraction.     

The spatial dynamics of crop and ground active predatory arthropods and their aphid prey in winter wheat

The distribution of aphid predators within arable fields has been previously examined using pitfall traps. With this technique predominantly larger invertebrate species are captured, especially Carabidae, but the technique provides no estimate of density unless mark-recapture is used. However, many other numerically important aphid predators occur in arable fields and relatively little is known about their distribution patterns nor whether they exhibit a density-dependent response to patches of cereal aphids. Identification of the most effective predators can allow management practices to be developed accordingly. In this study, the distribution of cereal aphids and their predators was examined by suction sampling within a field of winter wheat in Devon, UK, along with visual estimates of weed patchiness. Sampling was conducted on four occasions in 1999 across a grid of 128 sample locations. The distribution of 11 predatory taxa from the Carabidae, Staphylinidae and Linyphiidae was examined. Additionally, the total number of aphid predators and a predation index were used in these analyses. Carabid adults and larvae, along with staphylinid larvae showed the strongest aggregation into patches and the most temporal stability in their distribution. Other taxa had more ephemeral distributions as did the cereal aphids. The distribution of carabid larvae was disassociated from the distribution of cereal aphids for the first two sampling occasions indicating biocontrol was occurring. Other predatory groups showed both association and disassociation. Carabid larvae, Bathyphantes and total numbers of Linyphiidae showed a strong correlation with weed cover for two of the sample dates. Cereal aphids were disassociated from weed cover on three sampling occasions.     

The effect of weeds on the numbers of hoverfly (Diptera: Syrphidae) adults and the distribution and composition of their eggs in winter wheat

The abundance and species composition of syrphid adults in herbicide-treated and untreated headlands was recorded in a replicated within-field experiment conducted over a two-year period. The highest numbers of the most abundant syrphid species, Episyrphus balteatus, were recorded in the untreated headland strips. Analysis of the behaviour of the adult flies showed that they were retained in the untreated strips because they were foraging on the flowering non-crop plants. The distribution of syrphid eggs between herbicide treated and untreated headland replicates was examined. A significant positive relationship between the numbers of eggs per aphid and weed density was detected during June in the second year of the study. This is in part attributed to the tendency of Melanostoma females to oviposit on non-crop plants. The implications of the implementation of a modified herbicide regime in the headlands of cereal fields for the biological control of cereal aphids by syrphid larvae is discussed.     

Monoclonal antibodies reveal the potential of the tetragnathid spider Pachygnatha degeeri (Araneae: Tetragnathidae) as an aphid predator

The drive towards a more sustainable and integrated approach to pest management has engendered a renewed interest in conservation biological control, the role of natural enemy communities and their interactions with prey. Monoclonal antibodies have provided significant advances in enhancing our knowledge of trophic interactions and can be employed to help quantify predation on target species. The tetragnathid spider Pachygnatha degeeri Sundevall was collected from fields of winter wheat in the UK and assayed by ELISA for aphid proteins. It was demonstrated that this spider did not simply consume greater quantities of aphids because it was bigger. In addition, P. degeeri contained significantly greater concentrations of aphid in their guts than other spiders, showing that aphids comprised a greater proportion of their diet. Although P. degeeri constituted only 6% of the spider population numerically, females and males respectively contained 16% and 37% of total aphid proteins within all spiders screened, significantly more than their density would predict. These spiders also preyed upon aphids at a disproportionately high rate in June, during the aphid establishment phase, theoretically the best time for limiting growth in the aphid population. Although less abundant than other generalist predators, the capability of these hunting spiders to consume large numbers of aphids highlights them as a more significant component of the predator complex than had previously been realized. Limitation of aphid numbers early in the year by generalist predators provides more time for the specialist aphid predators and parasitoids to move in.     

The effect size of aphid‐tending ants in an agricultural tri‐trophic system

Most studies regarding ant–aphid interactions focus only on the direct effects of ants on tended aphids and aphidophagous predators, or the indirect effects on the host plant. Studies evaluating the effects of aphid‐tending ants on more than one trophic level are rare and evaluate only the presence or absence of such effects. Here we assessed the effect sizes of ants in a tri‐trophic system (common bean plants, aphids and lacewing larvae). We tested if the presence of aphid‐tending ants has positive effects on aphid abundance and host‐plant production and negative effects on aphid predator abundance. We also hypothesized that aphid‐tending ants affect more intensely trophic levels that are more directly related to them (i.e., first aphids, then aphid predators and then host plants). We tested these hypotheses in field mesocosms experiments using the presence and absence of ants. We found that aphid‐tending ants have great positive effects on final aphid abundance. Ants also positively affected the number of seeds; however, it was not possible to measure the effect size for this trophic level. Furthermore, ants had negative effects on lacewing larvae only at first release. The effect size of ants was greater for aphids, followed by lacewing larvae, and with no effects on the number of seeds produced. Ants positively affect aphids and host‐plant production, probably by way of honeydew collection preventing the development of entomophagous/saprophytic fungi. On the other hand, ants negatively affect lacewing larvae by excluding them from the host plant. In natural systems, several ant species may attend aphids, differently affecting the organisms of the various trophic levels within the ant–aphid interaction, thereby obscuring the real effect size of ants. Assessing the effect size of aphid‐tending ants on the organisms involved in ant–aphid interactions provides more realistic information about the effects of this interaction on natural systems.     

Ultralow oxygen treatment for postharvest control of Nasonovia ribisnigri (Homoptera: Aphididae) on iceberg lettuce

The aphid Nasonovia ribisnigri (Mosley) is a common pest of lettuce in the United States. It hinders export of U.S. lettuce to the overseas market such as Japan where it is a quarantined pest. Ultralow oxygen treatments were studied for control of the insect on iceberg lettuce. Small-scale ultralow oxygen treatments in plastic jars were conducted at 1, 5, and 10 degrees C for different durations to determine effective treatment against nymphs and alates of N. ribisnigri. At oxygen levels of 0.015-0.025%, N. ribisnigri can be controlled in 3 d at 1 degrees C, 2 d at 5 degrees C, and 1 d at 10 degrees C. Large-scale ultralow oxygen treatments were conducted in bulk container treatment chambers with commercial iceberg lettuce heads for 2 d at 6 degrees C with oxygen levels of 0.015 and 0.025% and for 3 d at 3 degrees C with oxygen level of 0.015%. All treatments achieved complete control of N. ribisnigri. No negative impact on lettuce quality was detected after 2 wk of posttreatment storage. Therefore, the selected treatments have potential to be commercially developed for postharvest control of N. ribisnigri on iceberg lettuce.