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.     

Larvae of the green lacewing <Emphasis Type="Italic">Mallada desjardinsi</Emphasis> (Neuroptera: Chrysopidae) protect themselves against aphid-tending ants by carrying dead aphids on their backs

Larvae of the green lacewing Mallada desjardinsi Navas are known to place dead aphids on their backs. To clarify the protective role of the carried dead aphids against ants and the advantages of carrying them for lacewing larvae on ant-tended aphid colonies, we carried out some laboratory experiments. In experiments that exposed lacewing larvae to ants, approximately 40% of the larvae without dead aphids were killed by ants, whereas no larvae carrying dead aphids were killed. The presence of the dead aphids did not affect the attack frequency of the ants. When we introduced the lacewing larvae onto plants colonized by ant-tended aphids, larvae with dead aphids stayed for longer on the plants and preyed on more aphids than larvae without dead aphids. Furthermore, the lacewing larvae with dead aphids were attacked less by ants than larvae without dead aphids. It is suggested that the presence of the dead aphids provides physical protection and attenuates ant aggression toward lacewing larvae on ant-tended aphid colonies.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Effect of spatial heterogeneity on the role of Coccinella septempunctata as an intra-guild predator of the aphid pathogen Pandora neoaphidis

The foraging behavior of starved and non-starved adult and larval Coccinella septempunctata on groups of plants in the presence of Pandora neoaphidis-infected Acyrthosiphon pisum, uninfected aphids or a mixture of these two prey types was compared. In general results of these studies confirmed the results of previous work comparing foraging behavior on a smaller spatial scale in Petri dishes. However, behaviors were modified in response to spatial complexity, prey quality, and the host plant. Starved C. septempunctata adults and larvae fed for longer and consumed more aphids than non-starved coccinellids. Both larvae and adults fed on infected aphids and in some cases entirely consumed them. This was thought to be due to the ease of capture of infected (dead) aphids and the feeding stimuli provided by the presence of the host plant and, where there was a choice of prey, uninfected aphids in the environment. Both larvae and adults spent the majority of the time foraging in the upper regions of plants and visited more plants when they were not starved or when they were in the presence of less suitable, infected aphid prey.     

Effectiveness of larvae ofSyrphus ribesii andS. Corollae [Diptera: Syrphidae] as predators onMyzus persicae [Homoptera: Aphididae]

Capture efficiency, handling time and functonal response to prey density were studied in larvae ofSyrphus ribesii (L.) andS. corrollae (Fabr.) eatingMyzus persicae Sulz. at 20°C, 16 hrs light. First instar larvae ofS. ribesii had distinctly higher capture efficiency than 1st instar larvae ofS. corollae, both versus 1st instar and adult aphids. Second and 3rd instar larvae of both species seemed to prefer adult rather than 1st instar aphids, but no distinct difference in capture efficiency between the species was found. On comparable stages,S. ribesii always had shorter handling time thanS. corollae and it appeared in both species to be correlated with size of prey and predator. Handling time was thus shortest when 3rd instar larvae consumed 1st instar aphids. During one hour, 3rd instar larvae ofS. ribesii consumed aphids in quantities almost linearly dependent on aphid density (5, 10, 20 and 40 adult aphids/100 cm²), although the response also could roughly be described byHolling's “basic functional response curve”. On the contrary, 3rd instar larvae ofS. corollae soon reached a maximum consumption during one hour, being almost constant (5–7 aphids) at prey densities ≥10 aphids/100 cm².

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Résumé L'efficacité prédatrice (proportion de toutes les rencontres entre larves de syrphes et pucerons se terminant par la capture et la succion de pucerons), la durée d'activité et la relation entre l'absorption de nourriture et la densité des proies (5, 10, 20 et 40Myzus persicae Sulz. aptères au 5^e stade larvaire/100 cm²) ont été étudiées chez des larves deSyrphus ribesii (L.) etS. corollae (Fabr.) à 20°C et 16 h de photopériode. Les L₁ deS. ribesii présentent une efficacité prédatrice des pucerons des premier et dernier stades nettement plus élevée que les L₁ deS. corollae (S. ribesii: 71 et 38%;S. corollae: 42 et 0%). Chez les larves des 2^e et 3^e stades, on n'a observé aucune différence d'efficacité prédatrice entre les 2 espèces, mais une préférence pour les pucerons adultes. La durée d'activité deS. ribesii est dans tous les cas plus courte que celle deS. corollae pour un même stade de larves et de pucerons. Une corrélation positive semble exister entre la taille des proies et celle des prédateurs. La durée d'activité est la plus courte pour des prédateurs en L₃ s'alimentant de pucerons au premier stade (1,3 mn chezS. ribesii et 2,3 mn chezS. corollae). L'absorption de nourriture enregistrée en une heure chezS. ribesii varie presque linéairement avec la densité des pucerons; elle correspond d'autre part assez bien à la formule deHolling. L'absorption de nourriture parS. corollae atteint un maximum de 7 pucerons à l'heure et demeure pratiquement constante à partir d'une densité≥10 pucerons/100 cm².

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The author wishes to thank techn. ass.Bodil Horgen for assistance during the experiments     

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.     

Behavioral Response to Larval Tracks and the Influence of Tracks on Intraguild Scavenging by Coccinellid Larvae

A paired design was used to determine that Harmonia axyridis 4th instars were not influenced by the presence of conspecific larval tracks, but well-fed H. axyridis 4th instars spent less time on plants that contained tracks left by Coleomegilla maculata 4th instars. To determine if the presence of larval tracks influences intraguild scavenging by H. axyridis 4th instars, dead 4th instars were placed in Petri dishes that contained or did not contain larval tracks. The presence of larval tracks did not influence the feeding frequency or the amount of time before feeding. However, larvae dragged their pygopod on dish surfaces more frequently if the dish contained larval tracks. In addition, starved H. axyridis larvae were more likely to feed on the prey and dragged their pygopod less frequently than well-fed larvae.     

Effect of prey availability on foraging and production efficiencies of larvalMetasyrphus corollae [Dipt.: Syrphidae]

We determined the effects of number of prey and the area over which they were distributed on the number of prey eaten and the production efficiency ofMetasyrphus corollae (F.) (formelySyrphus corollae F.) larvae. We hypothesized that, for a given number of prey offered per day, increased plant size would reduce foraging efficiency and production efficiency. Also, increased prey number was expected to increase production efficiency. Individual larvae were offered 10, 20, 40 or 60 pea aphids each day on plants with 7.5, 35 or 73 cm² surface area. When more aphids were offered, larvae ofM. corollae killed more of them but left larger aphid carcasses. Foraging efficiency (percentage of available aphids eaten) declined as aphid abundance and plant size increased. Foraging costs were not decreased by increasing prey density and therefore production efficiency was not increased.

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Résumé Les effets du nombre de proies et de la surface sur laquelle elles ont été distribuées, sur le nombre de proies mangées et sur le rendement de la production des larves deMetasyrphus corollae (F.) ont été déterminés. Nous avons supposé que, pour un nombre donné de proies offertes par jour, le rendement de la quête alimentaire et de la production diminueraient avec l'augmentation de la taille de la plante. De même, on supposait que le nombre croissant de proies augmentait le rendement de la production. On offrait chaque jour à des larves individuelles 10, 20, 40 ou 50 aphides du pois, sur des plantes dont la surface était 7,5, 35 ou 73 cm². Quand on offrait plus d'aphides, les larves deM. corollae les tuaient, mais elles laissaient alors de plus gros cadavres. Le rendement de la quête alimentaire (pourcentage d'aphides disponibles mangés) diminuait lorsque l'abondance des aphides et la taille de la plante augmentaient. Les couts de la quête alimentaire n'étaient pas réduits par l'augmentation du nombre de proies et ainsi le rendement de la production n'était pas augmenté.

     

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.     

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.     

Feeding economy of larvae of a flower fly,Metasyrphus corollae [Dip.: Syrphidae]: Partial consumption of prey

Metasyrphus corollae (F.) larvae ingested aphid contents at a decreasing rate over time because the contents of the aphid became increasingly difficult to obtain as the body of the prey was emptied. Starved larvae usually handled prey longer than wellfed larvae. Younger starved larvae fed longer than older larvae. Larvae starved for 24 h ate the most of a prey. Larvae held prey, on average, until 71% of dry mass was extracted after which the prey carcass was discarded. Prey handling time and amount of each prey consumed were determined by size and hunger of larvae and degree of depletion of prey contents.

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Résumé Les larves deMetasyrphus corollae (F.) ingèrent de plus en plus lentement le contenu des pucerons en fonction du temps car ce contenu est de plus en plus difficile à extraire puisque la proie se vide. Les larves qui ont été privées de nourriture se nourrissent généralement plus longuement que les larves bien nourries, et les jeunes larves affamées plus longuement que les plus agées. Les larves qui n'ont pas mangé depuis 24 h sont celles qui consomment le plus. Les larves retiennent la proie, en moyenne, jusqu'à ce que 71% de son poids sec ait été extrait, après quoi la carcasse de la proie est jetée. Le temps de manipulation et la quantité consommée de la proie sont déterminés par la taille et la faim de la larve, ainsi que par l'épuisement du contenu de la proie. Il n'y a aucune preuve consistante que les larves deM. corollae répondent directement au nombre de proies.

     

Fitness of the hover flies Episyrphus balteatus and Eupeodes corollae faced with limited larval prey

Differences in adult oviposition behavior of the aphidophagous hover flies Episyrphus balteatus (De Geer) and Eupeodes corollae (F.) (Diptera: Syrphidae) led us to formulate and test hypotheses concerning their larval behaviour. In laboratory experiments, larval E. balteatus lived longer when starved than larval E. corollae, and this difference increased with age at which starvation commenced. Larval E. balteatus crawled faster than larval E. corollae, and this difference increased under starvation. Limited aphid supply reduced larval and pupal survival more for E. corollae than for E. balteatus. However, the effect of limited aphid supply on adult body size did not differ for the two species. E. balteatus oviposition and larval behavior make it a more promising candidate for introducing to control Diuraphis noxia (Mordwilko) (Homoptera: Aphididae) where this aphid has become an exotic pest.     

Influence of aphid honeydew on the foraging behaviour of Hippodamia convergens larvae

1. Environmental cues associated with prey are known to increase predator foraging efficiency. Ladybird larvae are major predators of aphids. The sugary excretion of aphids (honeydew) has been proposed to serve as a prey‐associated cue for ladybird larvae. 2. Ladybird larvae are regularly found on the ground moving between plants or after falling off plants. The use of prey‐associated cues would be particularly beneficial for ladybird larvae on the ground in that such cues would help them to decide which plants to climb because aphids are patchily distributed within as well as amongst plants and, as a result, many plants are either not infested with aphids or do not host an aphid species of high nutritional value for ladybird larvae. 3. Laboratory experiments with larvae of Hippodamia convergens Guérin‐Méneville (Coleoptera: Coccinellidae) were carried out to explore whether honeydew accumulated on the ground is used as a foraging cue. The study also investigated whether, if honeydew is a foraging cue, larvae show differential responses to honeydew of high‐quality prey Acyrthosiphon pisum Harris compared with that of low‐quality prey Aphis fabae Scopoli (both: Homoptera: Aphididae). 4. Hippodamia convergens larvae stayed longer in areas containing honeydew but did not engage in longer bouts of searching. Furthermore, larvae did not distinguish between honeydew from high‐ and low‐quality aphid prey.     

The use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae)

Abstract.  1. To clarify the use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L., searching behaviour of ladybird larvae for Aphis craccivora Koch and Acyrthosiphon pisum Harris and the abundance of honeydew under aphid colonies were examined in laboratory experiments.
2. More larvae responded by climbing the plants with aphids than responded to plants without aphids. When the plants were replaced with sticks, in order to exclude visual and olfactory cues from plants and aphids, more larvae of C. septempunctata climbed sticks above the area that contained honeydew than climbed sticks above the area that did not contain honeydew. Then, ladybird larvae use honeydew as a contact kairomone when foraging for aphids.
3.  Aphis craccivora deposited a larger number of honeydew droplets beneath the plants than did similar numbers of A. pisum. Thus, C. septempunctata larvae licked more frequently the honeydew of A. craccivora than that of A. pisum and spent longer searching on the area containing honeydew of A. craccivora than that of A. pisum . Consequently, a larger number of larvae climbed a stick above honeydew of A. craccivora than that of A. pisum.
4. It may be also considered that C. septempunctata larvae can distinguish honeydew of the two aphid species and respond more strongly to A. craccivora than A. pisum.     

Laboratory study of cannibalism and interspecific predation in ladybirds

Abstract.

• 1 In the absence of aphids, adult females of Adalia bipunctata (L.) showed a greater reluctance to eat eggs than males.
• 2 Eggs and young larvae were more vulnerable to cannibalism than older larvae and starved larvae were more vulnerable than well-fed larvae.
• 3 Both egg and larval cannibalism is inversely related to the abundance of aphids.
• 4 Eggs are a better food, in terms of larval growth and survival, than aphids.
• 5 In the absence of aphids interspecific predation occurred, but not equally, between the coccinellids A.bipunctata, A.decempunctata (L.), Coccinella septempunctata L. and C.undecempunctata L.
• 6 Larvae and adults of A. bipunctata and C.septempunctata were reluctant to eat conspecific eggs painted with a water extract of the other species' eggs and larvae of C. septempunctata were more likely to die after eating a few eggs of A.bipunctata than vice versa.
• 7 These results indicate that cannibalism occurs mainly when aphid prey is scarce and is adaptive in that it improves the chances of survival, and coccinellids, to varying degrees, are defended against interspecific predation.

     

Feeding behaviour of the ladybeetle,Pseudoscymnus kurohime

A laboratory study was made of the feeding behaviour of the ladybeetlePseudoscymnus kurohime (Miyatake) when attacking the sugar cane woolly aphidCeratovacuna lanigera Zehntner. The 1st-instar ladybeetle larva was smaller than the 1st instar aphid nymph. All larval stages of the ladybeetle sucked out the body fluids of aphids and left their emptied corpses. The 1st, 2nd, and 3rd instar ladybeetle larvae mostly attacked 1st instar aphids, whereas the 4th-instar ladybeetle larvae attacked all stages of aphids. Ladybeetle adults ate mostly 1st-instar aphids. Young larvae attacked aphids in several different ways: (1) They crawled under an aphid, seized it by its underside and lifted it up. (2) They attacked new born nymphs at birth or shortly afterwards. (3) They fed on an aphid that had been captured by an older larva. The larvae preferred to seize with their mandibles the head or thorax of an aphid, while adults seized their prey by the abdomen. When attacked by an adult, 82% of the aphids secreted droplets from their abdominal cornicles, whereas only 7.2–12% secreted droplets when attacked by larvae. The 4th instar larvae more voracious than the younger larvae.     

Effect of a co-occurring aphid on the susceptibility of the Russian wheat aphid to lacewing predators

Previous field experiments indicated that the presence of the bird cherry-oat aphid, Rhopalosiphum padi (L.), on perennial grasses can decrease the effectiveness of predatory lacewings, Chrysoperla plorabunda (Fitch), in reducing populations of the Russian wheat aphid, Diuraphis noxia (Mordvilko). We tested the hypothesis that R. padi deflects predation away from D. noxia because it feeds in sites that are more accessible to predators. We quantified the behavior of lacewing larvae on crested wheatgrass plants bearing either D. noxia alone or an equal mixture of D. noxia and R. padi. On non-flowering plants, R. padi typically occurred on leaf sheaths or open blades, and was encountered and captured more often than D. noxia, which usually fed within immature, rolled leaves. Overall time-budgets of lacewings did not differ between the pure-D. noxia and mixed-species treatments, but >75% of the time spent consuming aphids in the mixed-species treatment was devoted to R. padi. On flowering plants, D. noxia usually aggregated on the flag leaf below the inflorescence, whereas R. padi occurred mostly on leaf sheaths. Predators again captured R. padi more often than D. noxia, and spent more time consuming aphids in the mixed-species treatment than in the pure-D. noxia treatment. These behavioral observations support the hypothesis that non-target prey can hamper the short-term effectiveness of biological-control agents against D. noxia.     

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.     

Role of syrphid larvae and other predators in suppressing aphid infestations in organic lettuce on California's Central Coast

Organic lettuce, Lactuca sativa L., growers on the Central Coast of California rely on conservation biological control to manage Nasonovia ribisnigri Mosley (Hemiptera: Aphididae) and other aphid pests of lettuce. In 2006, we carried out five replicated field trials to determine the importance of syrphid larvae in the suppression of N. ribisnigri and other aphids infesting organic romaine lettuce. We used Entrust, a spinosad-based insecticide approved for use on organic farms, to suppress syrphid larvae in aphid-infested romaine. Romaine treated with Entrust was unmarketable at harvest because of aphid infestation, whereas insecticide-free romaine was marketable. Syrphid larvae composed 85% or more of total predators in most trials, and they were the only predators consistently recovered from romaine that was infested with aphids early and largely aphid-free by harvest. The species mix of nonsyrphid predators varied from site to site. Applications of Entrust suppressed nonsyrphid predators in two trials, and so was an imperfect tool for selectively suppressing syrphid larvae. The relative importance of syrphid larvae and other predators in the conservation biological control of aphids in organic romaine is discussed. We conclude that syrphid larvae are primarily responsible for the suppression of aphids in organic romaine on California's Central Coast.