Predation byCheilomenes vicina [Coleoptera: Coccinellidae] on the cowpea aphid,Aphis craccivora [Homoptera: Aphididae]: Effect of prey stage and density

Consumption of larvae and females ofAphis craccivora Koch by 1st and 4th larvae and adults ofCheilomenes vicina (Muls.) was studied under fluctuating temperature (24–30°C). The early aphid instars were consumed in significantly greater numbers than later instars and females. The feeding rates ofC. vicina were significantly positively correlated with the population density of prey. The number of prey consumed daily by each predator stage tested, increased more steeply at lower than at higher prey densities, exhibiting thus the type 2 functional response

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Résumé Consommation des larves et des ♀ d'Aphis craccivora par les11^e stades, les 4^e stades et les adultes deCheilomenes vicina a été étudiée à une température variant de 24 à 30°C. Les jeunes stades du puceron sont consommés en nombre plus grand significativement que les derniers stades et les ♂. Les taux d'alimentation deC. vicina manifestent une corrélation hautement significative avec la densité de population de la proie. Le nombre de proies consommées quotidiennement par chaque stade considéré du prédateur augmentait plus brusquement aux faibles qu'aux fortes densités de proies, manifestant ainsi chez le prédateur le type 2 de réponse fonctionnelle.

     

Interactions among vegetable-infesting aphids,the fungal pathogen Metarhizium anisopliae (Ascomycota: Hypocreales) and the predatory coccinellid Cheilomenes lunata (Coleoptera: Coccinellidae)

Entomopathogenic fungi are among biocontrol agents being considered for the control of aphids on a variety of crops. Predatory coccinellids, although generalist, are also among important natural enemies that must be conserved for aphid management. Laboratory studies were carried out to investigate the interaction between three vegetable-infesting aphids, Metarhizium anisopliae isolate ICIPE 62 and the coccinellid predator Cheilomenes lunata. At a concentration of 1?×?10⁸?conidial?ml^–1, the fungus was found to cause mortality of 7.5% to C. lunata, compared to 2.5% mortality in the control at 10?days post-treatment. Female adult C. lunata to which fungus-infected aphids were offered as prey never accepted them as food source in non-choice bioassays. However, live and dead non-infected aphids were fed upon. In choice bioassay, a total of 1–3 out of 24 infected non-sporulating aphids per species (average of 0.1–0.4 aphids per arena) were consumed by 48?h-starved C. lunata within a period of 60?min, but avoided sporulating cadavers. Foraging adult C. lunata enhanced the spread of conidia of M. anisopliae from infected cadavers to fourth instars Aphis gossypii feeding on okra (0.8–15.0% mortality), Brevicoryne brassicae (3.3–15.0% mortality) and Lipaphis pseudobrassicae (0.8–14.2% mortality) on kale plants. Results of this study demonstrate compatibility between M. anisopliae and C. lunata, and could provide a sustainable strategy for effective management of aphids on crucifers and okra cropping systems.     

Evidence for utilization of Diptera in the diet of field-collected coccinellid larvae from an antibody-based detection system

Aphidophagous coccinellid larvae have a wide range of potential prey in alfalfa and during times of low aphid abundance, larvae may supplement their diet with alternative prey. To understand the effects of the seasonal aphid availability on alternative prey use, an order-specific monoclonal antibody, DrosW-VI-B8, was used to examine the frequency of dipteran predation by these important natural enemies. Over 400 larvae were hand-collected from alfalfa and, in parallel, arthropod abundance was recorded. Harmonia axyridis and Coccinella septempunctata larvae were abundant early in the season when aphid populations were at their peak and Coleomegilla maculata larvae were collected later in the season when potato leafhoppers were abundant in the alfalfa. A relatively low proportion of field-collected H. axyridis, C. septempunctata, and C. maculata tested positive for dipteran proteins throughout the season. Similar to prior studies examining stage differences in coccinellid food breadth, older instars tested positive for dipteran proteins (3rd instar, 6% positive; 4th instar, 7% positive) but no early instars screened positive. This study provides a valuable insight into the trophic linkages that exist between coccinellid larvae and Diptera.     

Relative toxicity of three wheat herbicides to two species of Coccinellidae

Abstract On the High Plains of the USA, herbicides specific for broad-leaf weeds are regularly applied to winter wheat in the early spring, sometimes late enough to coincide with the colonization of fields by cereal aphids and their natural enemies. We tested the toxicity of three such herbicides, Ally (Dupont), Rave (Syngenta) and 2,4-D ester (generic), to neonate larvae of two coccinellid species important in cereal aphid biocontrol, Coleomegilla maculata DeGeer and Hippodamia convergens Guérin-Méneville. Topical treatment of larvae with 2,4-D resulted in 25% and 60% mortality in the two species, respectively, with surviving C. maculata larvae experiencing a 5% increase in developmental time. No significant effects were noted for the other two materials, save for a 2.5% increase in developmental time for C. maculata larvae exposed to Rave. No material caused significant mortality in either species when larvae were fed on prey (Schizaphis graminum Rondani) treated with herbicide 24 h earlier, although 2,4-D reduced developmental time slightly in C. maculata. When herbicide applications are delayed enough in spring to coincide with aphid activity in wheat, farmers can reduce the risk of disrupting biological control by selecting an alternative to 2,4-D.     

The establishment ofRhyzobius forestieri [Col.: Coccinellidae] in greece and its efficiency as an auxiliary control agent against a heavy infestation ofSaissetia oleae [Hom.: Coccidae]

Introduction releases of the exotic coccinellid predatorRhyzobius forestieri Mulsant were made in 22 localities throughout Greece. The releases were made on olive, citrus, fig and almond trees infested with differentCoccidae scales. In 9 of these localities,R. forestieri was recovered 2–5 months later. The most important factor for the successful establishment ofR. forestieri seems to be an abundance of scales on the trees. To study the ability ofR. forestieri to enhance overall predation, 3 releases were made in an olive grove heavily infested withSaissetia oleae Olivier on Chios island during 1982 and 1983. The experiment began in June 1982. By April 1983, theS. oleae infestation was clearly in the process of being controlled by the coccinellid. A native coccinellid,Exochomus quadripustulatus L., contributed little during this period because of its obligatory estival diapause. The other main coccinellid,Chilocorus bipustulatus L., made an appreciable contribution toward achieving control during the summers of 1982 and 1983, but it was affected from July onward by increasing parasitism of its larvae. Both species are quiescent during the winter months. The introduced speciesR. forestieri was readily established in the olive grove and multiplied quickly. One month after the initial release,R. forestieri larvae made up most of the coccinellid larval population in the field. Thereafter until April 1983,R. forestieri larvae and adults comprised the dominant coccinellid population in the olive grove. During the winter months,R. forestieri remains active as long as temperature exceeds 8 °C. The important increase inR. forestieri's population size between July 1982 and April 1983 coincided with the increased rate of predation onS. oleae. This suggests thatS. forestieri played a key role in bringing theS. oleae infestation under control during this period. After April 1983, however, theR. forestieri population diminished rapidly andC. bipustulatus became the dominant coccinellid in the olive grove. It is believed that scarcity of prey was the cause of this reversal: by April 1983, the density ofS. oleae in the olive grove had already been reduced by about 100 times.     

Predation on Mindarus abietinus infestingbalsam fir grown as Christmas trees: the impact ofcoccinellid larval predation with emphasis on Anatis mali

The impact of natural coccinellid larvalpredation on the balsam twig aphid was evaluated bysystematically removing coccinellid egg masses in a6–8 year-old balsam fir (Abies balsamea)Christmas tree plantation in southwesternQuebec. Among coccinellid species hunting on firfoliage during development of Mindarus abietinusfundatrices in May, the indigenous Anatis mali was by far the most abundant and themain one to oviposit on trees. Comparison of trees onwhich coccinellid larval predation was excluded withcontrol trees showed that A. mali had a markedimpact both during and after the phase of rapid M. abietinus population growth that followedfundatrix maturation. On trees where coccinellidlarvae were allowed, aphid colonies became inactive(i.e. no live aphids in the colony) about two weeksearlier than on controls. A strong dampening effect onaphid density was also observed in those colonies thatremained active until the end of the aphid life cycle.Predation on aphid colonies reduced sexualsproduction, as the density of M. abietinusoverwintering eggs per shoot subsequently was reducedby 32%. Predation by coccinellid larvae occurred toolate to prevent needle damage to current year shoots,which affects the aesthetic value of Christmas trees.However, current year shoots measured in the mid-crownof trees late in the season were 19% longer on treeswhere aphid predation by coccinellid larvae wasallowed, compared with trees where they were excluded.Rearing all larval stages of A. mali on 4thinstar and adult sexuparae of M. abietinusindicated an average consumption of 269 aphids tocomplete larval development and pupate, which wasequivalent to at least seven colonies of M.abietinus at maximum aphid density at theexperimental site. Anatis mali is an importantnatural control factor of balsam twig aphid inChristmas tree plantations, hence its activity shouldbe protected and possibly stimulated by favourablepest management practices.     

Maintenance of ladybeetle diversity across a heterogeneous African agricultural/savanna land mosaic

The importance of agricultural land for maintaining indigenous biodiversity is an important contemporary issue. A large agricultural estate in Swaziland, which is a mosaic of citrus orchards, exotic tree windbreaks, indigenous riparian vegetation and savanna was investigated. The distribution of larval and adult coccinellids within the habitat types, and the relative influences of temperature, relative humidity, ground insolation and prey presence was assessed over a 2-year period. The highest coccinellid densities and species richness occurred within the orchards. Riparian habitats on the boundaries of the estate were closer to the orchard habitat in species composition than was the natural savanna. Not surprisingly, the presence/absence of prey was the most important variable which determined coccinellid distribution and density in the agricultural landscape. Relative humidity, temperature and ground insolation had no significant effect on coccinellid densities. Some species, such as the exotic Cheilomenes lunata, and the indigenous Scynmus sp. and Nephus sp. were widely dispersed and occurred in all habitats, while the economically important but exotic species, Rodolia cardinalis was only within the citrus orchards. Overall, the coccinellid assemblage was not a good indicator of habitat type. Nevertheless, a variety of habitat types (i.e. a heterogenous landscape) appear to be essential for the survival of most coccinellids. Indeed, the heterogeneous agricultural and natural land mosaic provides maximum plant diversity and hence coccinellid diversity, thus increasing the potential for natural control of pest prey species in the orchards, while at the same time maximizing survival of indigenous coccinellids.     

Coccinellid immigration to infested host patches influences suppression of Aphis glycines in soybean

Generalist natural enemies may be well adapted to annual crop systems in which pests and natural enemies re-colonize fields each year. In addition, for patchily-distributed pests, a natural enemy must disperse within a crop field to arrive at infested host patches. As they typically have longer generation times than their prey, theory suggests that generalist natural enemies need high immigration rates to and within fields to effectively suppress pest populations. The soybean aphid, Aphis glycines Matsumura, is a pest of an annual crop and is predominantly controlled by coccinellids. To test if rates of coccinellid arrival at aphid-infested patches are crucial for soybean aphid control, we experimentally varied coccinellid immigration to 1 m² soybean patches using selective barriers and measured effects on A. glycines populations. In a year with low ambient aphid pressure, naturally-occurring levels of coccinellid immigration to host patches were sufficient to suppress aphid populations, while decreasing coccinellid immigration rates resulted in large increases in soybean aphid populations within infested patches. Activity of other predators was low in this year, suggesting that most of the differences in aphid population growth were due to changes in coccinellid immigration. Alternatively, in a year in which alate aphids continually colonized plots, aphid suppression was incomplete and increased activity of other predatory taxa contributed to adult coccinellid predation of A. glycines. Our results suggest that in a system in which natural enemy populations cannot track pest populations through reproduction, immigration of natural enemies to infested patches can compensate and result in pest control.     

Cannibalism and interspecific predation in two predatory ladybirds in relation to prey abundance in the field

Life tables for two predatory ladybirds,Coccinella septempunctata andHarmonia axyridis, were constructed in two years in which prey abundance differed. The prey aphid,Aphis gossypii, was abundant on its primary host from late May to late June. Females ofH. axyridis andC. septempumctata laid their eggs during the increase and peak in aphid abundance. The oviposition period ofC. septempunctata started before and was shorter than that ofH. axyridis. In both species, intraguild predation and/or cannibalism of the egg and fourth instar stages, but rarely of other developmental stages, were observed. Fourth instar larvae ofH. axyridis had to complete their development when aphids were scarce more frequently than those ofC. septempunctata. The highest frequency of intra- and interspecific predation was of fourth instarH. axyridis larvae. In terms of intraguild predation, the larvae ofH. axyridis preyed on larvae ofC. septempunctata, but the reverse was not observed. Percentage survival from egg to adult inH. axyridis was higher than inC. septempunctata in both years, and least in both species in the year of low aphid abundance. These results suggest that prey abundance influenced the frequency of cannibalism and intraguild predation, which were important in structuring this guild of ladybirds.     

Cornicle secretions of Uroleucon nigrotuberculatum (Homoptera: Aphididae) as the last bullet against lady beetle larvae

Aphids have evolved various defense strategies against natural enemies, including secretions from their cornicles. We assessed the defensive function of cornicle secretions by the goldenrod aphid, Uroleucon nigrotuberculatum (Olive), against larvae of the lady beetles Coccinella septempunctata bruckii Mulsant and Propylea japonica (Thunberg) (Coleoptera: Coccinellidae). The aphid secreted red droplets from its cornicles when attacked by the larvae. Two‐thirds of the C. septempunctata bruckii larvae and 46.7% of the P. japonica larvae that preyed on the aphids died before reaching the pre‐pupal stage. The secretions caused molting failure when smeared on the larvae's heads or glued to the larvae's mouthparts, killing 56.7% of C. septempunctata bruckii larvae and 36.7% of P. japonica larvae. Second instar larvae were affected most. About 40% of third and fourth instar larvae of C. septempunctata bruckii vomited soon after ingesting the aphids. In the field, up to 40% of first and second instar larvae were smeared with red secretions. Our results show that these cornicle secretions are an effective and active defense against earlier instars of coccinellid larvae.     

Stage and age-specific predation in four aphidophagous ladybird beetles

In the present study, stage- and age-specific predatory patterns and nutritional ecology attributes of four co-occurring aphidophagous ladybirds, namely, Coccinella septempunctata (L.), Coccinella transversalis F., Cheilomenes sexmaculata (F.) and Propylea dissecta (Mulsant) on pea aphid, Acyrthosiphon pisum (Harris) were investigated. Despite their lower consumption rates, smaller ladybirds (C. sexmaculata and P. dissecta) and earlier instars were efficient converters of aphid prey with higher relative growth rates, over the respective larger ladybirds (C. septempunctata and C. transversalis) and fourth instars. The consumption rates of pea aphids by the larvae increased with stage, but with a decline in the later part of the fourth instars, just prior to pupation. Adult females of early age groups (between 2 and 7 days) had higher consumption rates than later age groups in all the four species. Further in adult females, consumption rates decrease with increase in age, but without significant changes in conversion efficiencies and relative growth rates.     

Intraguild responses of aphid predators on apple to the invasion of an exotic species, Harmonia axyridis

The effects of the invasion ofan exotic predator, Harmonia axyridis(Pallas) (Coleoptera: Coccinellidae), wereinvestigated using three experiments on theecology of aphid predators on apple. Oneexperiment, 1992, was collected prior to theH. axyridis invasion, and two others,1996 to 1997 and 1999 to 2000, were collectedafter the invasion. Except for one year, 1999,H. axyridis was the dominant coccinellid,replacing the formerly dominant Coccinellaseptempunctata L. (Coleoptera: Coccinellidae),another exotic species. The dominance of H. axyridis was greater among larvae thanamong adults. There was no apparent effect ofthe H. axyridis invasion on abundance ofthe predator, Aphidoletes aphidimyza(Rondani) (Diptera: Cecidomyiidae) and apossible positive effect on the abundance ofchrysopids (Neuroptera: Chrysopidae). Principal component analysis indicated thatalthough individual species were affected, theoverall effect of H. axyridis invasion onthe predator guild as a whole was negligible. The data indicate that the interaction betweenthe two exotic species, H. axyridis andC. septempunctata, may be allowing nativecoccinellids to become more abundant on applethan when C. septempunctata was thedominant coccinellid.     

Deterrent effects of larval tracks on conspecific larvae in <Emphasis Type="Italic">Cycloneda limbifer</Emphasis>

The response of Cycloneda limbifer Say (Coleoptera: Coccinellidae) larvae to conspecific larval tracks was investigated with the assistance of the automatic video tracking system EthoVision. Single larvae were monitored for 1 h in an arena with a choice of two substrates, one clean and one contaminated with the tracks of conspecific larvae. Fourth instar larvae of C. limbifer effectively differentiated clean substrates from substrates with fresh tracks of conspecific first instars. Larvae walked significantly further on clean than on contaminated substrates in each 20-min period of the 1 h trials. With the exception of the first 20-min period, residence times were also longer on clean than on contaminated substrates. The mean velocity of larvae on the substrates did not differ in any period of the test. The results indicate that a deterring pheromone in fresh tracks of C. limbifer larvae effectively deters conspecific larvae from staying on sites previously exposed to larvae. The mode of action of the semiochemical repellent secreted by larvae is complementary to a previously described semiochemical-mediated egg-spacing mechanism in the coccinellid C. limbifer. Active distribution of larvae has the potential to improve the initial distribution of offspring by females among aphid colonies of varying qualities and may facilitate survival during changes in the distribution of conspecific larvae in the course of the development.     

Adult Coccinellid Activity and Predation on Aphids in Spring Cereals

Improved understanding of coccinellid activity and predation on aphids in the field could clarify their potential in aphid biological control. Our objective was to determine the influence of abiotic and biotic factors on activity and predation by adults of three coccinellid species (Hippodamia convergens Guerin-Meneville, H. tredecimpunctata tibialis (Say), and Coleomegilla maculata DeGeer) in spring cereal fields. The proportion of time coccinellids spent searching was correlated with air temperature, aphid density, and time of day. The relationship between searching and these variables differed among species. Mean walking speed ranged from 66.4 cm/min for C. maculata to 83.2 cm/min for H. tredecimpunctata and increased with temperature for all species. The frequency of short flights (<2 m) by beetles increased with temperature and decreased with increasing aphid density for all species, but the relationship of short flight frequency to these variables differed among species. The frequency of long flights (>2 m) was similar for all species and influenced by temperature and calendar date. The frequencies with which aphids were encountered and eaten were correlated with aphid density and temperature for H. convergens and H. tredecimpunctata. Despite over 250 h spent observing adult coccinellid behavior in the field, predation data were insufficient to develop a useful predation model.     

Potential for augmentative biological control of black bean aphid in California sugarbeet

In northern California, black bean aphid (Aphis fabae Scopoli complex) can be a major pest of sugarbeet, particularly in spring-planted fields. The major natural enemies of the aphid are predators, including the coccinellidsHippodamia convergens Guerin,Coccinella novemnotata Herbst &C. septempunctata L., and the chrysopidsChrysopa oculata Say &C. nigricornis Burmeister. Augmentative releases of eggs of eitherChrysoperla carnea (Stephens) orC. rufilabris (Burmeister) failed to significantly reduce aphid populations under field conditions. This was attributed in part to incompatibility between black bean aphids and lacewing larvae from commercial sources. Application of food sprays containing yeast hydrolysate, sucrose and molasses resulted in increased densities of eggs and adults ofC. carnea in treated plots. These results suggest that the potential for augmentative biological control of black bean aphid through the application of food sprays is greater than that for release of commercially available lacewings. Regardless of method, an holistic approach to augmentation that takes into account the ecological structure of the target agroecosystem will be required. Some aspects of “pre-emptive” biological control are discussed.     

Developmental Changes in the Sterol Composition and the Glycerol Content of Cuticular and Internal Lipids of Three Species of Flies

The glycerol concentration and the composition of cuticular and internal sterols in three medically and forensically important fly species, viz., Musca domestica, Sarcophaga carnaria, and Calliphora vicina, were analyzed. The cuticular and internal lipid extracts were separated by HPLC‐LLSD, after which the sterol fraction was characterized by GC/MS in total ion current (TIC) mode. The cuticular lipids of M. domestica larvae contained seven sterols, while in pupae and females, six sterols were identified. Five sterols were found in the cuticular lipids of M. domestica males. The internal lipids of M. domestica larvae and pupae contained six and seven sterols, respectively, while those of male and female flies contained only five sterols. Sitosterol, cholesterol, and campesterol were the dominant sterols in M. domestica, while campestanol, stigmasterol, sitostanol, and fucosterol were identified in low concentrations or in traces. In contrast, cuticular and internal lipids of S. carnaria and C. vicina contained only cholesterol. Glycerol was identified in all stages of M. domestica, S. carnaria, and C. vicina. For all the three examined fly species, the present study clearly showed species‐specific developmental changes in the composition of cuticular and internal sterols as well as in the glycerol concentration.     

Asymmetric competition between larvae of the blowflies Calliphora vicina and Lucilia sericata in carrion

1. The relative abundance of the blowflies Calliphora vicina (R.-D.) and Lucilia sericata (Meigen) in carrion was considered in relation to inter- and intraspecific larval competition and the distribution of adults between habitat types. 2. In mixed and pure laboratory cultures of L. sericata and C. vicina the mortality of both species increased and adult size declined as the initial larval number was increased. However, for L. sericata at initial numbers greater than ten larvae/g of liver, the effects of competition on adult size and mortality were greater in the mixed cultures than in the pure cultures. In contrast, for C. vicina at numbers greater than ten larvae/g of liver, the effects of competition on adult size and mortality were greater in the pure cultures than in the mixed cultures. 3. The laboratory data suggest therefore that for L. sericata the effects of interspecific competition with C. vicina on size and mortality were greater than the effects of intraspecific competition in a pure culture at the same initial number. Notably, however, the intensity of interspecific competition was not sufficiently asymmetric to allow C. vicina to exclude L. sericata even at the highest numbers examined. 4. In the field, higher numbers of adult L. sericata emerged from the carcasses of laboratory mice placed in open pasture than in woodland or hedgerow sites. In contrast, higher numbers of C. vicina emerged from carcasses placed in woodland and hedgerow sites. 5. Trapping showed that in the field adult L. sericata were relatively more abundant in open pasture than in woodland and hedgerow sites, while C. vicina were more abundant in woodland and hedgerow sites than in open pasture. 6. It is concluded that the low numbers of L. sericata that emerge from carrion relative to the numbers of C. vicina may, in part, be the result of asymmetric interspecific competition, but that the uneven distribution of adults of the two species among habitat types also plays a major role in shaping the blowfly community in carrion.     

Effects of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae), on three non-target coccinellids

A microsporidium from Hippodamia convergens was transmitted horizontally to three non-target coccinellid hosts (Adalia bipunctata L., Coccinella septempunctata L. and Harmonia axyridis Pallas) under laboratory conditions. For all species examined, microsporidia-infected larvae took significantly longer to develop than did uninfected larvae but the microsporidium had no effect on larval mortality. Adult sex ratios of uninfected and microsporidia-infected adults were about 1:1 (♀:♂) and did not differ significantly. At the end of a 90-day trial, microsporidia-infected H. convergens produced significantly fewer eggs and did not live as long as uninfected individuals. Differences in fecundity and longevity were not observed for the three non-target coccinellids that were examined. Mean spore counts from smear preparations of microsporidia-infected A. bipunctata did not differ significantly from H. convergens, suggesting that A. bipunctata (a native coccinellid) is a suitable host for the microsporidium but infection was lighter in C. septempunctata and H. axyridis (introduced species). Vertical transmission of the pathogen was observed during the 90-day trial by examining eggs and larvae that were produced by microsporidia-infected adults. For all species examined, 100% vertical transmission of the pathogen was eventually observed. Three eugregarines were found in two adult A. bipunctata: Gregarine A trophozoites are similar in size to those of Gregarina katherina Watson (described earlier from Coccinella spp.), Gregarine B trophozoites are similar in size to those of Gregarine A but are morphology distinct, and Gregarine C trophozoites are similar in size to G. barbarara Watson (described earlier from A. bipunctata).     

Instar-specific defense ofColeomegilla maculata lengi (Col.: Coccinellidae): Influence on attack success of the intraguild predatorChrysoperla rufilabris (Neur.: Chrysopidae)

We conducted a laboratory study of the behavioural interactions between larval stages of the lady beetleColeomegilla maculata lengi Timberlake and an intraguild predator, the lacewingChrysoperla rufilabris Burmeister. Ladybird susceptibility to predation by third instarC. rufilabris is age-specific and influenced by larval defensive strategies. First and second instars ofC. maculata were captured more consistently by chrysopid larvae than third and fourth instars. Young coccinellids generally exhibit escape reactions (dropping, fleeing, retreating) but, do not survive after being caught by lacewings. In addition to escape reactions, older larvae may actively defend themselves by wriggling or biting, even after being pierced. The proportion of physical interactions that resulted in coccinellid death was significantly lower for older instars (<2%) than for first (35%) and second (19%) instars. Capture efficiency byC. rufilabris was related to the coccinellid-chrysopid size ratio. Dropping off the leaves, the most common defensive behaviour displayed by coccinellid larvae, contributed to reduce overall mortality of first instars to a level similar to the one observed for older stages. However, lacewings were able to prevent coccinellid dropping by impaling and holding their prey in the air, or by falling with the ladybirds to the ground where they continued their attack.