Influence of cornicle droplet secretions of the cabbage aphid,Brevicoryne brassicae,on parasitism behavior of na＇fve and experienced Diaeretiella rapae

Insects have evolved amazing methods of defense to ward off enemies. Many aphids release cornicle secretions when attacked by predators and parasitoids. These se cretions contain an alarm pheromone that alerts other colony members of danger, thereby providing indirect fitness benefits to the releaser. In addition, contact with cornicle se cretions could also threaten an attacker and could provide direct fitness to the releaser. However, cornicle secretions may also be recruited as a kairomonal cue by aphid natural enemies. In this study, we investigated the effect of the cornicle droplet volatiles of the cabbage aphid, Brevicoryne brassicae （L.）, on the hostsearching behavior of naive and experienced female Diaeretiella rapae （M＇Intosh） parasitoids in olfactometer studies. In addition, we evaluated the role ofB. brassicae cornicle droplets on the oviposition prefer ence of the parasitoid in a twochoice bioassay. Naive females did not exhibit any preference between volatiles from aphids secreting cornicle droplets over nonsecreting aphids, while experienced parasitoids exploited the secretions in their host location. Experienced females were also able to choose volatiles from both secreting and nonsecreting aphids over clean air, while this ability was not observed in naive females. Although secretion of cornicle droplets did not influence the percentage of first attack in either naive or experienced females, the success of attack （i.e. resulting in a larva） was significantly different between secreting and nonsecreting aphids in the case of experienced parasitoids.     

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.     

Body size affects host defensive behavior and progeny fitness in a parasitoid wasp,Lysiphlebus fabarum

This study tested effects of maternal body size on foraging behavior and progeny development in a thelytokous population of Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). Small and large wasps were reared from first and second instar hosts [black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae)], respectively, and each was provided with a patch (bean leaf disk) containing either 15 small (second instar) or 15 large (fourth instar) hosts for a 30‐min foraging period. Neither body size nor host size affected time allocation to various behaviors within a patch, but second instar aphids produced significantly more mummies than fourth instars. The preferred attack orientation was from the side of the aphid, suggesting wasps were sensitive to the risk of smearing with cornicle secretions. Few wasps developed in fourth instar hosts, suggesting later host instars were somewhat resistant to parasitism. Second instar hosts, the most suitable stage for L. fabarum development, relied more on defensive behavior, specifically kicking and secreting cornicle droplets. Large wasps were more likely to elicit a double cornicle secretion, indicating that aphids graded their response to the size of the attacker. Larger wasps were also more likely to be smeared with cornicle secretion, suggesting they were more vulnerable than small wasps. Although small wasps had smaller eggs than large wasps, there was no effect of maternal egg size on the size of progeny. However, daughters of small females emerged with larger egg loads than daughters of large mothers, and their eggs tended to be slightly smaller, although not significantly. Regression analysis revealed a positive correlation between maternal egg size and progeny developmental time for small and large wasps, and between maternal egg size and progeny egg load for small wasps. These results confirm maternal effects of body size in an aphid parasitoid, and reveal that vulnerability to host behavioral defenses is also body size dependent.     

三种瓢虫集团内捕食卵对初龄幼虫生长发育的影响

种内和种间捕食卵现象常见于对食蚜瓢虫的研究报道中。源自亚洲、入侵北美和欧洲的异色瓢虫(Harmonia axyridis)由于卵含有生物碱而被认为可能是威胁本土瓢虫生存的原因,但该推测只有在明确异色瓢虫在原产地是否对本土瓢虫具有类似负面影响后,才能确证。通过饲喂实验,旨在明确异色瓢虫在原产地是否对本土七星瓢虫(Coccinella septempunctata)和龟纹瓢虫(Propylea japonica)具有食卵负面作用,为揭示异色瓢虫入侵机理提供依据。实验中用蚜虫(对照)和3种瓢虫卵对瓢虫初孵1龄幼虫进行饲喂处理,观察幼虫生长发育指标。对瓢虫初龄幼虫存活表现的分析结果表明:(1)龟纹瓢虫初孵幼虫取食同种卵和七星瓢虫卵后存活至2龄的比率均高于取食异色瓢虫卵的近7倍;(2)七星瓢虫幼虫取食异色瓢虫卵未存活到2龄,但取食其他瓢虫卵和蚜虫后85%以上个体存活至2龄;(3)异色瓢虫初孵幼虫取食同种和其他2种瓢虫卵后有90%以上存活至2龄。龟纹瓢虫和七星瓢虫1龄幼虫取食异种卵比同种卵后的发育历期显著延长,但异色瓢虫并未表现出差异。龟纹瓢虫和异色瓢虫1龄幼虫取食同种和异种卵后的体增重相同,但七星瓢虫取食龟纹瓢虫卵后体增重比取食同种卵减小。研究结果说明,异色瓢虫卵对七星瓢虫和龟纹瓢虫初孵幼虫的生存具有负面影响,反之不然。所以,取食异色瓢虫卵本身并非是导致北美和欧洲本土瓢虫数量减少的原因。     

不同虫态及虫龄大草蛉与七星瓢虫之间的集团内互作

【目的】为明确新疆棉田棉蚜 Aphis gossypii 捕食性天敌之间的集团内捕食效应及其对蚜虫数量的控制作用。【方法】本研究以优势天敌昆虫大草蛉 Chrysopa pallens 和七星瓢虫 Coccinella septempunctata 为对象,以棉蚜为猎物,在温室中利用盆栽棉花,首先观察了2种天敌昆虫之间各虫态及虫龄配对的19个处理在无蚜植株上共存24 h后的存活数,然后观察了2种天敌昆虫配对处理下棉苗上棉蚜数量随时间的变化趋势。【结果】在无蚜棉株上2种捕食性天敌昆虫共存24 h后的存活结果表明：（1）在发育阶段相同的配对组合中,若是成虫则均存活,若是１龄幼虫则大草蛉存活较多,若是末龄幼虫则七星瓢虫存活较多;（2）在有卵的组配中,除七星瓢虫卵不被大草蛉成虫所捕食外,其他5个组配处理中卵均被捕食;（3）在有蛹的配对组合中,除七星瓢虫蛹被大草蛉末龄幼虫捕食外,其他处理下蛹均不被捕食;（4）在成虫与幼虫的配对组合中,七星瓢虫成虫捕食较多的大草蛉1龄幼虫,但不捕食大草蛉末龄幼虫,而大草蛉成虫与七星瓢虫1龄或末龄幼虫之间不发生捕食;(5）在不同龄期幼虫的配对组合中,大草蛉末龄幼虫捕食七星瓢虫1龄幼虫,而七星瓢虫末龄幼虫捕食大草蛉1龄幼虫。在有蚜植株上2种捕食性天敌共存对棉蚜数量具有不同的控制作用：（1）2种捕食昆虫的幼虫各自单独存在（对照）下,蚜虫密度随时间而降低;（2）大草蛉幼虫与七星瓢虫幼虫或成虫配对处理下,棉蚜密度随时间而增大;（3）大草蛉成虫与七星瓢虫幼虫或成虫配对处理下,棉蚜密度随时间而减小。【结论】研究结果说明,大草蛉与七星瓢虫之间存在集团内捕食,但２种天敌共存对棉蚜的控制作用取决于大草蛉虫态, 若大草蛉为幼虫,可使蚜虫密度增大,若为成虫,则使蚜虫密度减小。     

Effects of neem-treated aphids as food/hosts on their predators and parasitoids

Abstract:  Experiments were conducted in the laboratory and in greenhouses. Of three neem preparations sprayed upon eggs, only neem oil (NO) exerted a negative impact on the hatching rate of Coccinella septempunctata and Chrysoperla carnea . First instar larvae of Episyrphus balteatus proved to be highly susceptible, when feeding 24 h on aphids sprayed with neem kernel water extract (NKWE). First instar larvae of C. septempunctata showed a very high mortality when feeding on aphids sprayed with different neem preparations. Aphid feeding and live span was reduced. When NKWE had been applied to the soil, the mortality of larvae of E. balteatus and C. septempunctata were lower, when feeding on aphids. Second instar larvae of C. septempunctata were far less susceptible when feeding 48 h on neem-sprayed aphids than first instars; the time of their development was prolonged, and aphid consumption reduced. Larvae of C. carnea proved to be less susceptible, when feeding on neem-sprayed aphids, than E. balteatus and C. septempunctata . In C. carnea , however, significant influences were also observed in aphid consumption, time of development, mortality, longevity, and rate of deformity. NO, containing a very low concentration of azadirachtin A, had stronger negative effects than NeemAzal-T/S^®, in all observations. In the parasitoid Diaeretiella rapae , NKWE application to the soil induced negative reactions, when aphids on these plants were parasitized: low percentage parasitization, lowered mummy weight, low emergence rate of adults of F1 and even of F2. Foliar sprays of NKWE had less severe effects in this parasitoid species. The results are discussed with regard to their theoretical and practical significance.     

A comparison of triglycerides from aphids and their cornicle secretions

Direct mass spectrometry of extracts showed that body triglycerides from 30 species of aphids contained the same fatty acid radicals, C₆ (hexanoyl), C_6:2 (sorboyl), C₁₄ (myristoyl), and C₁₆ (palmitoyl) as did the cornicle secretions, but in many species the proportions of hexanoyl and/or palmitoyl triglycerides were greater in the body. When cornicle secretions were collected progressively so as to draw increasingly upon body fat reserves, their composition changed gradually towards that of the body extracts.All summer forms of Myzus persicae had similar body triglycerides, even when selected for resistance to organophosphorus insecticides, or bred for 3 months on an artificial diet. The composition of body triglycerides was also independent of colour in two aphid species in which pink and green forms were compared.Body extracts contain enough triglycerides for their composition to be determined in single aphids and the use of body extracts allows examination of aphids lacking cornicles and of specimens that do not give cornicle secretion because of low body turgor. Although, as in the case of cornicle secretions, the triglyceride composition of body extracts was not well correlated with taxonomic position, body extracts provide a second chemical characteristic that can be used to define a particular species.     

Cornicle length in Macrosiphini aphids: a comparison of ecological traits

Abstract 1. Aphids often emit cornicle droplets when attacked by predators. While the function of cornicle droplets has long been debated (i.e. mechanical protection vs. chemical signalling), it is not understood why aphid species have cornicles of different lengths.
2. It was hypothesised that aphids living in more scattered colonies have longer cornicles to scent-mark predators with cornicle droplets containing alarm pheromone, so that clone-mates are provided with advanced warning of a threat, even if not at the predation site. To test this hypothesis, multiple regression analyses were used, due to a lack of phylogenetic information on these taxa, to address which ecological traits (amount of wax on an aphid, degree of colony aggregation, feeding shelter, ant attendance) are correlated with cornicle length.
3. Aphids living in dense colonies tended to have shorter cornicles than aphids living in more scattered colonies. Also, aphids with more protection (i.e. wax) on their bodies had shorter cornicles. Aphids also tended to have shorter cornicles when tended by ants. The presence of a feeding shelter was not a good predictor of cornicle length.
4. It is suggested that longer cornicles function to scent-mark predators with alarm pheromone to increase the inclusive fitness of a clone; however the negative correlation between the amount of individual protection, and also ant attendance, and cornicle length argues for a trade-off between different forms of defence.     

Relative consumption of three aphid species by the lacewing, Chrysoperla rufilabris, and effects on its development and survival

Relative consumption of three aphid species, Aphis gossypii Glover, Myzus persicae (Sulzer) and Lipaphis erysimi (Kaltenbach) (Homoptera: Aphididae), by larvae of the lacewing, Chrysoperla rufilabris (Burmeister) (Neuroptera: Chrysopidae), was determined in the laboratory, together with effects on lacewing development and survival. Percentages of survival of C. rufilabris from first instar to adult eclosion were significantly different among lacewing larvae fed different aphid species. When larvae were fed A. gossypii and M. persicae, all larvae developed to adulthood. All larvae died prematurely when they were fed L. erysimi. Developmental duration of C. rufilabris larvae was significantly shorter when larvae were fed A. gossypii (18.0 d) than when larvae were fed M. persicae (19.2 d). The number of fourth instar aphids consumed during development by C. rufilabris larvae differed significantly among individuals fed different aphid species. Chrysoperla rufilabris consumed an average of 168 M. persicae, followed by 141.6 A. gossypii, and only 26.6 L. erysimi. The percentage of these total number of aphids consumed by each larval stadium of C. rufilabris varied significantly among aphid species. The percentage of A. gossypii consumed by each larval stadium was similar to that for M. persicae, 12.1 and 11.4% by the first instar, 15.7 and 13.1% by the second instar, and 72.2 and 75.5% by the third instar, respectively; whereas in the case of L. erysimi, 23.3% of the total number of aphids were consumed by the first instar, 30.1% by the second instar, and 46.6% by the third instar.     

Soybean Aphid Response to their Alarm Pheromone E-ß-Farnesene (EBF)

When attacked by natural enemies some insect pests, including many aphid species, alert neighboring conspecifics with alarm pheromones. Cornicle secretions with pheromones benefit the attacked aphid but are costly to produce, while alarm pheromone benefits probably fall largely on alerted conspecifics. Given these variable benefits, the likelihood of a secretion may change depending on aphid density. Thus, we first hypothesized that the common alarm pheromone in aphids, E-ß-farnesene (EBF), was present in soybean aphid (Aphis glycines Matsumura) cornicle secretions and would elicit an alarm response in aphids exposed to it. Second, since aphids other than the secretor also benefit from cornicle secretions, we hypothesized that the likelihood of secretion would increase concurrently with the density of neighboring clonal conspecifics. Third, because alarm reaction behavior (e.g. feeding cessation) is probably costly, we hypothesized that alarm reaction behavior would decrease as conspecific density (i.e. alternative prey for an attacking natural enemy) increased. We found that soybean aphids 1) produce cornicle secretions using EBF as an alarm pheromone, 2) are less likely to release cornicle secretions when alone than in a small group (~10 individuals), but that the rate of secretion does not increase further with additional conspecific density, and 3) also exhibit alarm reaction behavior in response to cornicle secretions independent of aphid density. We show that soybean aphids can use their cornicle secretions to warn their neighbors of probable attack by natural enemies, but that both secretion and alarm reaction behavior does not change as density of nearby conspecifics rises above a few individuals.     

Chemical composition of cornicle secretion of the brown citrus aphid Toxoptera citricida

Brown citrus aphid Toxoptera citricida Kirkadly is considered as an important pest of citrus because it vectors citrus tristeza closterovirus. Aphids secrete a fluid from their cornicles as a defensive mechanism against natural enemies. Earlier studies on cornicle secretions of aphids focus only on triglycerides and fatty acids. In the present study, three different methods are used to investigate the chemical composition of the cornicle fluid of T. citricida. Gas chromatography with flame ionization detection is used to detect and quantify the triglycerides after trimethylsilyl derivatization, and gas chromatography‐mass spectrometry (GC‐MS) is used to determine the fatty acid composition after derivatization with boron trifluoride–methanol. Other compounds are detected using GC‐MS after methoxyamine hydrochloride and N‐methyl‐N‐(trimethylsilyl)trifluoroacetamide derivatization. The major fatty acid in the cornicle secretion of T. citricida is palmitic acid. Oleic, stearic, myristic, myristoleic and sorbic acids are also detected, although in low amounts. Sorboyl, dipalmitoyl (C_6‐2, C₁₆, C₁₆) and disorboyl, stearoyl (C_6‐2, C_6‐2, C₁₈) are the main triglycerides detected in cornicle secretion. Trehalose is the most predominant sugar (558.2 mm ), followed by glucose (92.0 mm ) and inositol (48.8 mm ). Many amino acids, including proline, glycine, alanine and serine, are also detected. In addition, the cornicle secretion is rich in many organic acids, including malic, citric, succinic and lactic acid. Information obtained from the present study improves our understanding of the chemical composition of the cornicle secretion of the brown citrus aphid.     

Population expansion in the North African Late Pleistocene signalled by mitochondrial DNA haplogroup U6

Background

Altruistic anti-predatory behaviours pose an evolutionary problem because they are costly to the actor and beneficial to the recipients. Altruistic behaviours can evolve through indirect fitness benefits when directed toward kin. The altruistic nature of anti-predatory behaviours is often difficult to establish because the actor can obtain direct fitness benefits, or the behaviour could result from selfish coercion by others, especially in eusocial animals. Non-eusocial parthenogenetically reproducing aphids form colonies of clone-mates, which are ideal to test the altruistic nature of anti-predatory defence behaviours. Many aphids release cornicle secretions when attacked by natural enemies such as parasitoids. These secretions contain an alarm pheromone that alerts neighbours (clone-mates) of danger, thereby providing indirect fitness benefits to the actor. However, contact with cornicle secretions also hampers an attacker and could provide direct fitness to the actor.

Results

We tested the hypothesis that cornicle secretions are altruistic by assessing direct and indirect fitness consequences of smearing cornicle secretions onto an attacker, and by manipulating the number of clone-mates that could benefit from the behaviour. We observed parasitoids, Aphidius rhopalosiphi, foraging singly in patches of the cereal aphid Sitobion avenae of varied patch size (2, 6, and 12 aphids). Aphids that smeared parasitoids did not benefit from a reduced probability of parasitism, or increase the parasitoids' handling time. Smeared parasitoids, however, spent proportionately more time grooming and less time foraging, which resulted in a decreased host-encounter and oviposition rate within the host patch. In addition, individual smearing rate increased with the number of clone-mates in the colony.

Conclusions

Cornicle secretions of aphids were altruistic against parasitoids, as they provided no direct fitness benefits to secretion-releasing individuals, only indirect fitness benefits through neighbouring clone-mates. Moreover, the use of cornicle secretions was consistent with their altruistic nature, because the occurrence of this behaviour increased with the size of indirect fitness benefits, the number of clone-mates that can benefit. This study provides evidence for a case of kin-directed altruistic defence outside eusocial animals.     

Development and functional response of Coelophora inaequalis (Coleoptera: Coccinellidae) feeding on brown citrus aphid, Toxoptera citricida (Homoptera: Aphididae)

1 Development and survivorship of Coelophora inaequalis (F.) were evaluated in the laboratory on the brown citrus aphid, Toxoptera citricida (Kirkaldy) at 20 and 25 °C. Coelophora inaequalis could complete its life cycle feeding on T. citricida at these two temperatures. The developmental period from egg to adult was significantly longer at 20 °C (24.7 d) than at 25 °C (15.9 d). The survivorship of combined immature stages were higher at 25 °C (39.2%) than at 20 °C (34.3%). 2 The effects of prey densities (4–64 aphids per leaf disk) on the functional responses of C. inaequalis fourth‐instar larvae and female adults were investigated using grapefruit leaf disks over a 24‐h period at 25 °C and a photoperiod of LD 14 : 10 h. Logistic regression analyses indicated that fourth‐instar larvae and adults exhibited a type II functional response to aphid density. Fourth‐instar larvae had a higher search rate (0.1305) than C. inaequalis female adults (0.0989). The handling time of fourth‐instar larvae (30.4 min) was significantly longer than that of female adults (5.8 min). Collectively, C. inaequalis appears to be a promising biological control agent of T. citricida in the citrus groves.     

Adaptations of Coccinella magnifica Redtenbacher, a Myrmecophilous Coccinellid, to Aggression by Wood Ants (Formica rufa Group). I. Adult Behavioral Adaptation, Its Ecological Context and Evolution

Coccinella magnifica is an obligate associate of Formica rufa-group ants. The congener Coccinella septempunctata can serve as a model for its nonmyrmecophilous ancestor. Formica rufa behavior toward these two ladybirds, and their behavior, was compared. Although C. magnifica was rarely attacked on ant trails, it was usually attacked on tended aphid colonies. Coccinella septempunctata was more readily attacked. The two ladybirds' behavior was similar on trails, but C. magnifica used more defense and C. septempunctata more rapid escape behavior on aphid colonies. Only C. magnifica fed upon tended aphids. Chemical adaptation to overcome ant aggression probably exists in C. magnifica, but it possesses almost no novel behaviors to counter ant aggression. Instead, modifications have occurred in the expression of behaviors present in C. septempunctata.     

Direct vs. inclusive fitness in the evolution of aphid cornicle length

By comparing the relative sizes of anatomical structures among phenotypes, selective pressures that shape species' morphologies can be evaluated. Aphids emit droplets containing an alarm pheromone/defensive secretion from unique anatomical structures called cornicles, upon being attacked. As aphids live in colonies of high relatedness, it is uncertain whether direct or inclusive fitness benefits have chiefly promoted cornicle evolution. Morphological measurements for apterous parthenogen, alate parthenogen, female sexual and male sexual morphs of 43 species (21 genera, one subfamily) were assessed to distinguish between the hypotheses that: (1) cornicles evolved for mechanical defence against natural enemies (direct fitness); (2) cornicles evolved for alarm signalling (inclusive fitness); or (3) cornicle length has been largely constrained by flight aerodynamics. Our results generally support the inclusive fitness hypothesis; cornicle length decreases as the relative number and relatedness of offspring decreases. As cornicle length is greatest in apterous parthenogenetic morphs, inclusive fitness benefits of protecting highly related kin may have been a key factor selecting for cornicles, and increased cornicle length, in aphids.     

Behavioural responses of the seven‐spot ladybird Coccinella septempunctata to plant headspace chemicals collected from four crop Brassicas and Arabidopsis thaliana,infested with Myzus persicae

1 Insects using olfactory stimuli to forage for prey/hosts are proposed to encounter a ‘reliability–detectability problem’, where the usability of a stimulus depends on its reliability as an indicator of herbivore presence and its detectability. 2 We investigated this theory using the responses of female seven‐spot ladybirds Coccinella septempunctata (Coleoptera: Coccinellidae) to plant headspace chemicals collected from the peach‐potato aphid Myzus persicae and four commercially available Brassica cultivars; Brassica rapa L. cultivar ‘turnip purple top’, Brassica juncea L. cultivar ‘red giant mustard’, Brassica napus L. cultivar ‘Apex’, Brassica napus L. cultivar ‘Courage’ and Arabidopsis thaliana. For each cultivar/species, responses to plants that were undamaged, previously infested by M. persicae and infested with M. persicae, were investigated using dual‐choice Petri dish bioassays and circular arenas. 3 There was no evidence that ladybirds responded to headspace chemicals from aphids alone. Ladybirds significantly preferred headspace chemicals from B. napus cv. Apex that were undamaged compared with those from plants infested with aphids. For the other four species/cultivars, there was a consistent trend of the predators being recorded more often in the half of the Petri dish containing plant headspace chemicals from previously damaged and infested plants compared with those from undamaged ones. Furthermore, the mean distance ladybirds walked to reach aphid‐infested A. thaliana was significantly shorter than to reach undamaged plants. These results suggest that aphid‐induced plant chemicals could act as an arrestment or possibly an attractant stimulus to C. septempunctata. However, it is also possible that C. septempunctata could have been responding to aphid products, such as honeydew, transferred to the previously damaged and infested plants. 4 The results provide evidence to support the ‘reliability–detectability’ theory and suggest that the effectiveness of C. septempunctata as a natural enemy of aphids may be strongly affected by which species and cultivar of Brassica are being grown.     

Searching efficiency of the lady beetleCoccinella septempunctata larvae in uniform and patchy environments

Experiments were conducted on the searching behavior and searching efficiency of the lady beetleCoccinella septempunctata bruckii Mulsant under conditions of various prey distributions and prey densities. The larvae changed their searching behavior before and after feeding. Before feeding the larvae moved quickly and the searching paths were nearly linear. But after feeding the speed decreased and turning angle increased. The speed and turning angle reverted gradually and recovered the initial pattern 95 s after feeding. The searching efficiency differed depending on the prey distribution. At low prey density, searching was most efficient when prey were distributed uniformly. But at middle and high prey densities, searching was most efficient when prey items were highly aggregated. The observed searching behavior of 4th instarC. septempunctata larvae was likely to be optimal considering the natural distribution of colonies of their prey, aphids.     

The role of vision and color in the close proximity foraging behavior of four coccinellid species

The role of vision and color in close-proximity foraging behavior was investigated for four species of lady beetles: Coccinella septempunctata, Hippodamia convergens, Harmonia axyridis, and Coleomegilla maculata. The effect of light level and color cues on consumption rates varied among the four predator species. The consumption rates of these predators on the pea aphid Acyrthosiphon pisum (Harris) was measured under light and dark conditions. C. septempunctata,H. convergens, and Ha. axyridis consumed significantly more aphids in the light than in the dark, while the consumption rate of Col. maculata was not affected by light level. Foraging ability was also measured on red and green color morphs of the pea aphid on red, green, and white backgrounds. C. septempunctata consumed significantly more of the aphid morph that contrasted with the background color, and showed no difference between morphs on the white background. H. axyridis consumed significantly more red morph aphids regardless of background. The remaining two species showed no difference in consumption rates on the two color morphs. The variation in the use of visual cues demonstrates how different species of predators can exhibit different foraging behaviors when searching for the same prey. Received: 4 August 1997 / Accepted: 3 February 1998     

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.     

Predation capacity and prey preference of <Emphasis Type="Italic">Chrysoperla carnea</Emphasis> on <Emphasis Type="Italic">Pieris brassicae</Emphasis>

The predation capacity and prey preference of larvae of Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) on eggs or larvae of Pieris brassicae (Linnaeus) (Lepidoptera: Pieridae) in the absence and presence of cabbage aphids as an alternative prey were evaluated in laboratory experiments at 25°C. Both instars preyed upon butterfly eggs and larvae as well as on cabbage aphids with the third instar being the most voracious. The lacewings had a strong preference for caterpillars to butterfly eggs. In the presence of the aphids the predation on P. brassicae eggs or larvae was either completely abandoned or reduced by about 70%, respectively, by second instar lacewings and either reduced by about 80% or maintained, respectively, by third instar lacewings. Both instars thus had a clear preference for aphids compared to eggs of P. brassicae. However, second instar lacewings preferred aphids to caterpillars whereas the opposite was the case for third instar lacewings. The results indicate that 3rd instar C. carnea has a potential as biocontrol agent against P. brassicae.