Presence of two host races of Aphis gossypii Glover (Hemiptera: Aphididae) collected in Turkey

The aphid, Aphis gossypii, is a primary pest of citrus, cotton, cucurbits and greenhouse‐grown vegetables in Turkey and throughout Europe. There is some previous empirical data suggesting that host‐adapted genotypes of this aphid exist which may in fact be host‐races. To determine if host races of A. gossypii are indeed present in the eastern Mediterranean region of Turkey, reciprocal host transfer experiments and life table analyses were performed with multiple asexual lineages (= clones) of the aphid collected from different hosts. The collection hosts included citrus, cucumber, eggplant, okra, sweet pepper and cotton. Aphid developmental times on the host from which the aphid was originally collected (= collection or natal host) were shorter (5.2–6.0 days) and had a higher intrinsic rate of population growth (r_m = 0.25–0.44) than the 6.6–7.3 days required when the aphid was reared on a non‐original collection host (= non‐collection host or non‐natal host) and had r_m = 0.03–0.30. Total immature mortality of the cotton clone, especially in the first nymphal stage, was high (51–100%) with low r_m (0–0.03) on cucumber, citrus and sweet pepper. Aphid populations transferred from citrus, eggplant and okra to cotton (r_m = 0.29–0.30) did not differ significantly in their performance from that of the cotton population on cotton (r_m = 0.34), whereas that from sweet pepper and cucumber populations (r_m = 0.22–0.24) were significantly lower. These data have allowed us to separate A. gossypii into two distinct biological groups: (a) a ‘generalist’ population obtained from cucumber, sweet pepper, citrus, eggplant and okra which exhibited statistically better development on cotton; versus (b) a population from cotton which, by comparison on reciprocal hosts, developed poorly on non‐natal hosts except on eggplant. Development of the cotton clone on cucumber and okra was not improved after four successive generations on the non‐natal host. The good development of A. gossypii from eggplant and cotton on these reciprocal hosts suggests that these particular clones were similar, if not identical, host races.     

The role of plant chemical cues in determining host preference in the generalist aphid parasitoid Aphidius colemani

The host preference behaviour of the generalist aphid parasitoid Aphidius colemaniwas investigated using a Y-tube olfactometer. Female A. colemanishowed a preference for the host-plant complex on which they had been reared, even though the same aphid host was involved, demonstrating a host plant preference. This preference was not evident when the parasitoids were dissected from their mummies prior to adult emergence. Host plant preference exhibited during host selection appeared to be induced by chemical cues encountered on the mummy case at the time of emergence, but preferences could be changed by subsequent foraging experiences. It is concluded that plant chemical cues play a major role in determining initial preferences through a process of emergence conditioning but that learning processes, involving cues encountered during oviposition in or contact with the host, can modify these initial preferences.     

The effect of rearing history and aphid density on volatile‐mediated foraging behaviour of Diaeretiella rapae

1. Parasitoid females foraging for hosts rely on cues derived from the insect host, the host plant and/or their interaction, and all of these can be learned during the immature and adult stages. 2. The present study investigated the importance of rearing history on foraging behaviour of Diaeretiella rapae, an endoparasitoid often associated with aphids feeding on brassicaceous plant species. 3. Parasitoids were reared on one of the four possible combinations, comprising two brassicaceous host plant species, Brassica nigra or Raphanus sativus, and two aphid species Brevicoryne brassicae or Myzus persicae. These parasitoids were tested in a Y‐tube olfactometer and given the choice between volatiles emitted by an aphid‐infested plant (25 or 100 aphids per plant) and an uninfested control plant. The parasitoid's responses were compared when offered: (i) the same plant–aphid combination as the one on which it had been reared; (ii) the same host plant infested with the alternative aphid species; or (iii) an alternative plant with the alternative aphid species. 4. Aphid density did affect the behavioural responses to the various odour sources, but rearing history did not. Diaeretiella rapae only preferred aphid‐induced to non‐induced plant volatiles at low aphid infestation level, whereas they did not discriminate between volatiles at high aphid infestation level. 5. It is concluded that aphid‐induced volatiles of brassicaceous plants play an important role during host habitat location, but seem less important for parasitoids to locate the aphid host itself. The data are discussed in the light of manipulation of host plant defences by aphids.     

Host selection and parasitism behavior of Lysiphlebus testaceipes: Role of plant,aphid species and instar

The aphid parasitoid Lysiphlebus testaceipes is a potentially valuable biological control agent of Aphis gossypii a major worldwide pest of cotton. One means of increasing the abundance of a biological control agent is to provide an alternative host habitat adjacent to cropping, from which they can provide pest control services in the crop. Host selection and parasitism rate of an alternative host aphid, Aphis craccivora by L. testaceipes were studied in a series of experiments that tested its host suitability relative to A. gossypii on cotton, hibiscus and mungbean. Host acceptance, as measured by number of ovipositions was much greater in A. craccivora compared to A. gossypii, while more host aphids were accepted on mungbean than cotton. When given a choice L. testaceipes attacks more 4th instar and adult stages (63% and 70%, respectively) of both hosts than 2nd instar nymphs (47%). In a switching (host choice) experiment, L. testaceipes preferentially attacked A. craccivora on mungbean over A. gossypii on cotton. Observations of parasitoid contact with A. gossypii cornicle secretion suggest it provides a useful deterrent against parasitoid attack. From these experiments it appears L. testaceipes has a preference for A. craccivora and mungbean compared to A. gossypii and cotton, in this respect using A. craccivora and mungbean as alternative habitat may not work as the parasitoid is unlikely to switch away from its preferred host.     

Olfactory response of the anthocorid predatory bug Orius sauteri to thrips-infested eggplants

The predator Orius sauteri (Poppius) (Heteroptera: Anthocoridae) is an effective arthropod natural enemy of thrips, especially Thrips palmi Karny (Thysanoptera: Thripidae), a serious pest of vegetables. First, we studied O. sauteri's response to the odour from thrips-infested eggplant [Solamum melongena L. (Solanaceae)] in a laboratory by two-choice experiments using a Y-tube olfactometer. When detached eggplant leaves were used as odour sources, O. sauteri preferred the volatiles from uninfested leaves to clean air. Concerning preferences among differently infested leaves, O. sauteri preferred the volatiles from plants infested with 10–100 thrips per leaf to uninfested leaves, but showed no significant preference for artificially damaged leaves over uninfested leaves. Similar results were obtained when complete plants were tested as the odour source. Second, release and recapture experiments in a greenhouse, a more realistic set of conditions, were conducted to confirm whether a significant preference for infested plants occurred at similar infestation levels as in the laboratory. Trends favouring infested plants were detected at densities of five and 500 thrips per plant; however, at the five thrips per plant, this trend was due to the large deviation seen in infested plants in only one replicate. In light of the low tolerable thrips density of eggplant, it is necessary to confirm whether artificial treatments with chemicals induce the emission of herbivore-induced plant volatiles that can attract O. sauteri and prolong its residence time on the leaf.     

The influence of cultivar and cultivar-aphid odours on the olfactory response of the parasitoid Aphidius colemani

The olfactory response of the parasitoid Aphidius colemani (Viereck) (Hymenoptera: Braconidae) to odours in a tritrophic system involving three cultivars of common cabbage, Brassica oleracea var capitata, characterized by different levels of susceptibility to Myzus persicae (Sulzer) (Hemiptera: Aphididae) was studied in a four‐way olfactometer. Odours influenced A. colemani response in the olfactometer to varying degrees. The magnitude of parasitoid response to odours of uninfested cabbage depended on cultivar, with Derby Day [green‐leaved, susceptible to M. persicae and the crucifer specialist, Brevicoryne brassicae (Linnaeus) (Hemiptera: Aphididae)] and Minicole (green‐leaved, partially resistant with known antibiosis factors for B. brassicae) preferred over Ruby Ball (red‐leaved with antixenosis factors for M. persicae and B. brassicae). The odour of the cabbage cultivar on which the parasitoid had been reared was preferred over the other cultivars. However, when provided with a choice between odours of infested plants, parasitoids did not show a significant preference for the cultivar on which they were reared. Results from the study show that parasitioids differentiated between odour of the three cultivars in dependence of their rearing history when the plant is uninfested.     

Prey Habitat Location by the Cassava Mealybug Predator Exochomus flaviventris: Olfactory Responses to Odor of Plant, Mealybug, Plant–Mealybug Complex, and Plant–Mealybug–Natural Enemy Complex

Exochomus flaviventris Mader is considered to be the most active predator of the cassava mealybug Phenacoccus manihoti Matile–Ferrero in Central Africa. The response of experienced gravid female coccinellids to the odor of cassava plant (var. Zanaga), unparasitized mealybugs, plant–mealybug complex with or without feeding prey (parasitized or not), and plant–mealybug complex with or without conspecific coccinellids was investigated in a Y-tube olfactometer. The odor of uninfested cassava plants was not more attractive than clean air. Dual-choice tests revealed that mealybug-infested plants were preferred to mealybugs alone and mealybug-damaged plants and were the major sources of volatiles that attract females coccinellids to the microhabitat of its prey. The emission of volatile chemicals did not appear to be limited to the infested parts of the plant but did occur systemically throughout the plant. The presence of conspecific coccinellid larvae or adult males did not modify the attractiveness of the mealybug-infested plants. However, when an infested plant with conspecific predator females (alone or with conspecific males) was compared to an infested plant or infested plant with conspecific males, E. flaviventris females showed a preference for the last two sources of odor. The uninfested plant with conspecific males was also preferred to the uninfested plant with conspecific females. In addition, the odor of conspecific males was preferred over that of conspecific females. Female predators preferred the plant infested with unparasitized mealybugs over the plant infested with mealybugs previously parasitized. These results showed that E. flaviventris females use herbivore-induced plant volatiles during foraging and can detect via olfaction the presence of conspecific gravid females and parasitized prey, thus assessing patch suitability from a distance.     

Ant attendance of the cotton aphid is beneficial for okra plants: deciphering multitrophic interactions

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Effects of aphid honeydew sugars on the longevity and fecundity of the aphidophagous gall midge Aphidoletes aphidimyza

The predatory gall midge Aphidoletes aphidimyza, which is used for augmentative biological control of aphids in greenhouses, uses aphid honeydew as an energy source. Currently, Rhopalosiphum padi on barley plants and Melanaphis sacchari on sorghum plants are used as the banker plant systems for A. aphidimyza to control Aphis gossypii on eggplant in Japan. We analyzed the sugar components of the honeydew of these three species by HPLC. The major components of honeydew were sucrose, fructose, and melezitose for A. gossypii; glucose and fructose for R. padi; and glucose, fructose, and melezitose for M. sacchari. Maltose and trehalose were minor components of the honeydew of these species. We investigated the influence of sugars, including three artificial aphid honeydews, six sugar components, and distilled water, on the longevity of unmated A. aphidimyza. Both females and males attained the greatest longevity on sucrose and artificial honeydew of A. gossypii. Mean longevities of both females and males were shortest when they were provided only with water. We conducted another experiment in which a mated female was released in a cage with an eggplant seedling infested with A. gossypii and fed with sucrose or only water. Females fed with sucrose lived significantly longer and had higher lifetime fecundity than those fed only water.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

Antibiosis and antixenosis of chrysanthemum cultivars to the aphidAphis gossypii

An assessment was made of the antibiotic and antixenotic resistance of three cultivars of autumn flowering chrysanthemum (derived fromDendranthema morifolium (Ramat) Tzvelev andD. indicum (L.) Desmoulins) to the aphidAphis gossypii Glover. Antibiosis was also assessed at the top, middle and lower leaves of the plant. The three cultivars showed a range of resistance with Hero being the most susceptible, Surfine the most resistant and Purple Anne intermediate. There was a trend for antibiosis to improve with position down the plant, however there were cultivar by position interactions. Antixenosis was assessed at two levels, the first being under conditions where the aphid had complete access to the plant in caged choice assays and the second being under conditions where the aphid only had host odour to assess under olfactometer choice conditions. Preferences under the caged choice conditions found Hero to be the most preferred cultivar while there was no preference shown between Purple Anne and Surfine. However, olfactometer assessments showed Hero to be the least preferred of the three cultivars and, again, no preferences could be found between Purple Anne and Surfine.     

Adult experience modifies attraction of the leafminer parasitoidOpius dissitus (Hymenoptera: Braconidae) to volatile semiochemicals

Oviposition-experienced females of Opius dissitus Muesebeck, a braconid parasitoid of Liriomyza sativaeBlanchard, preferentially landed on leafminer-infested rather than uninfested lima bean (Phaseolus lunatus L.) plants in a flight tunnel assay. Both naive and oviposition-experiencedparasitoids responded strongly to odors of infested lima bean plants in a four-arm olfactometer in comparison with odors of uninfested plants, suggesting that volatile semiochemicals are used in host location. Parasitoids with an oviposition experience on lima bean (lima-experienced) spent significantly more time in the infested odor than naive individuals, however, eggplant-experienced wasps did not spend significantly more time in the infested odor field than naive wasps. When parasitoids reared on leafminers in lima bean were provided a choice between the odor of infested lima bean and the odor of infested eggplant or cotton, naive and lima-experienced wasps preferred infested lima odor. An oviposition experience on the other plant species resulted in a dramatic shift in preference. It was concluded that the experience effect was due, at least in part, to associative learning, as has been reported for other parasitoids. The parasitoids may perceive unconditioned stimuli during host contact and oviposition on an infested leaf and may associate those stimuli with volatile semiochemicals emanating from the leaf or host. Subsequently, the volatiles associated with the presence of hosts are used in directing the search for hosts.     

Specialized host-plant performance of the cotton aphid is altered by experience

Although distinct host specialization is observed for the cotton-melon aphid (Aphis gossypii Glover) on cotton and cucurbit plants, it is still ambiguous whether the specialization is altered by experience on a novel host plant. Here the performance of cotton and cucurbit-specialized aphids, A. gossypii on novel host plants was studied by a host-selection test and by the life-table method. The two host-specialized aphids cannot survive and establish populations after reciprocal host transfers. They have ability to recognize the host plants on which they were reared, and escape behavior from novel hosts was observed. Interestingly, the cotton and cucurbit-specialized aphids survive and reproduce normally on hibiscus (Hibiscus syriacus), a main overwintering host plant, and host-fidelity of A. gossypii to cucurbit plants is altered by feeding and living experience on hibiscus, which confers the same capacity to use cotton and cucumber on to the cucurbit-specialized population, but host-fidelity to cotton is not altered and the fitness of the cotton specialized population to cucumber is still poorer. A. gossypii from hibiscus has a significant preference for cotton to cucumber in the host-selection process, and none stays on cucumber more than 20 h after transfer. The results presented imply that cucurbit-specialized aphids might not return to an overwintering host plant (hibiscus) in wild fields, so host conservatism to cucurbit plants is maintained. The potential of cucurbit-specialized aphids of A. gossypii to use cotton plants, intermediated by experience on hibiscus, suggests that the specialized host-plant performance of phytophagous insects is not wholly conservative.     

Response of the soybean aphid parasitoid Binodoxys communis to olfactory cues from target and non-target host-plant complexes

The parasitoid Binodoxys communis (Gahan) (Hymenoptera: Braconidae) is a candidate for release against the Asian soybean aphid, Aphis glycines Matsumura (Homoptera: Aphididae), in North America. Using a series of Y-tube olfactometer assays, we recorded responses of naïve and experienced B. communis females to odors from both target and non-target host plant complexes. The non-target host plant complexes included three aphid species native to North America [Aphis monardae Oestlund, Aphis oestlundi Gillette, and Aphis asclepiadis (Fitch)] and one species presumed to be exotic to North America (Aphis nerii Boyer de Fonscolombe), all on their respective host plants. We also investigated whether the parasitoid distinguishes between volatiles from soybean infested with A. glycines vs. those of uninfested plants. Volatiles from A. glycines-infested soybean plants and several non-target host plant complexes elicited a response in B. communis, which seemed to be reinforced through oviposition experience. Wasps with experience on a specific host plant complex increased their response to odors of this complex. Neither naïve nor experienced wasps, however, preferred odors from target vs. non-target host plant complexes in choice tests. Our data also indicate that B. communis is unable to distinguish between volatiles from infested vs. uninfested soybean plants. This study shows that B. communis females respond to a broad array of olfactory stimuli, exhibit low fidelity for any particular odor, and employ some behavioral plasticity in their response to volatile cues. We discuss implications of these results for establishment of B. communis and potential undesirable non-target effects associated with the release of this species in North America.     

The role of chemical cues in host finding and acceptance by Callosobruchus chinensis

We studied the response of female Callosobruchus chinensis to chemical cues emitted by cowpea seeds at different stages of bruchid infestation (uninfested, egg carrying, L1-, and L4-infested). Olfactory attractiveness was determined in Y-tube olfactometer assays by testing individual seed categories against either clean air or uninfested seeds. Oviposition preferences between uninfested and infested seeds were determined in petri-dish choice-experiments. The olfactometer assays revealed that weevils discriminate between seeds containing different stages of developing bruchids on the basis of olfactory cues. While odors from uninfested and egg-carrying seeds acted as attractants, odors from L1- and L4-infested seeds failed to induce a positive response by the bruchids. When given a choice between uninfested and infested seeds in the olfactometer, weevils preferred uninfested seeds over L1- and L4-infested seeds, but failed to distinguish between uninfested and egg-carrying seeds. In the oviposition experiment as well, bruchids showed a distinct preference for uninfested seeds when offered in combination with L1- and L4-infested seeds. This experiment further showed a reduced acceptance of egg carrying seeds. Our results indicate that C. chinensis females use chemical information during both host searching and host acceptance. Volatiles from uninfested or egg carrying seeds act as attractants, while deterrence increases as development of bruchid immature stages progresses.     

Anthocorid predators learn to associate herbivore-induced plant volatiles with presence or absence of prey

We investigated how the plant‐inhabiting, anthocorid predator, Anthocoris nemoralis, copes with variation in prey, host plant and associated herbivore‐induced plant volatiles and in particular whether the preference for these plant odours is innate or acquired. We found a marked difference between the olfactory response of orchard‐caught predators and that of their first generation reared on flour moth eggs in the laboratory, i.e. under conditions free of herbivory‐induced volatiles. Whereas the orchard‐caught predators preferred odour from psyllid‐infested pear leaves, when offered against clean air in a Y‐tube olfactometer, the laboratory‐reared first generation of (naive) predators did not. The same difference was found when a single component (methyl salicylate) of the herbivore‐induced plant volatiles was offered against clean air. After experiencing methyl salicylate with prey, however, the laboratory‐reared predators showed a pronounced preference for this volatile. This acquired preference did not depend on whether the volatile had been experienced in the juvenile period or in the adult phase, but it did depend on whether it had been offered in presence or absence of prey. In the first case, they were attracted to the plant volatile in subsequent olfactometer experiments, but when the volatile had been offered during a period of prey deprivation, the predators were not attracted. We conclude that associative learning is the most likely mechanism underlying acquired odour preference.     

Singing on the wings! Male wing fanning performances affect female willingness to copulate in the aphid parasitoid Lysiphlebus testaceipes (Hymenoptera: Braconidae: Aphidiinae)

Lysiphlebus testaceipes (Hymenoptera: Braconidae: Aphidiinae) is a generalist endoparasitoid attacking more than 100 aphid species. In L. testaceipes, wing fanning is a main male courtship display evoked by a female‐borne sex pheromone. However, no information is available on the characteristics and behavioral role of male fanning during courtship in this parasitoid. Here, the courtship behavior of a wild strain of L. testaceipes was quantified and the male wing fanning performances were analyzed through high‐speed video recordings and examined in relation to mating success. Courtship sequence of wild L. testaceipes did not substantially differ from that previously reported for other populations mass reared on aphids. We observed that the male courtship duration did not affect mating success. However, video analysis revealed that the males producing high‐frequency fanning signals achieved higher mating success over those that display low‐frequency fanning. Wing fanning before successful and unsuccessful courtship differed in amplitude of wing movements and alignment toward the mate, highlighting that frontal courtship positively influence the female mating decisions. This study increases knowledge on sexual behavior in a key parasitoid of aphids, highlighting the importance of wing fanning among the range of sensory modalities used in the sexual communication of L. testaceipes. From a practical point of view, this information is useful in L. testaceipes‐based biocontrol strategies, since it can help to establish parameters for quality checking of mass‐reared wasps over time.     

Host instar suitability of Aphis gossypii (Hemiptera: Aphididae) for Lysiphlebus testaceipes (Hymenoptera: Braconidae) and parasitism effect on aphid life table

Lysiphlebus testaceipes (Cresson) (Hymenoptera: Braconidae, Aphidiinae) has constituted a well-studied parasitoid insect model, but very little is known about the host-instar suitability of aphid for the wasp so far. One of the hosts of L. testaceipes is Aphis gossypii Glover (Hemiptera: Aphididae). The latter is a serious aphid pest to vegetable production in Benin. Therefore, the objectives of our study were to: (1) examine the oviposition behavior of L. testaceipes on A. gossypii; (2) investigate the host-instar suitability of A. gossypii for L. testaceipes; and (3) compare the life table parameters of A. gossypii with aphids parasitized by L. testaceipes and unparasitized aphids (control). The study was conducted in a laboratory at 26 ± 1 °C in petri dishes and revealed that the parasitoid utilized up to seven stabbing stings to handle and oviposit, particularly in older A. gossypii. In aphids parasitized at the third instar, the net reproductive rate R _o as well as the intrinsic rate of natural increase r _m was significantly lower (2.119 ± 0.272 and 0.110 ± 0.018) compared to the control (15.529 ± 1.287 and 0.272 ± 0.008), respectively (p < 0.01).     

Olfactory responses of Aphidius gifuensis to odors of host plants and aphid‐plant complexes

Abstract The olfactory responses of Aphidius gifuensis to odors from two host plants (Nicotiana tabacum and Brassica napus ssp.) and their complexes with different infestation levels of two host aphids (Myzus persicae and Lipaphis erysimi) were respectively examined in an olfactometer. The results showed that female A. gifuensis did not respond to odors of undamaged or mechanically damaged host plants, but significantly responded to odors of aphid/plant complexes. Moreover, A. gifuensis responded significantly to odors of both M. persicae and L. erysimi/plant complexes when host plants were infested by high levels of aphids, suggesting that quantity of aphid‐induced volatiles could be important for attracting A. gifuensis. When tested between aphid/plant complexes, A. gifuensis did not show its preference for either complex. The efficiency of A. gifuensis against aphids in open fields potentially could be improved by using its olfactory response to aphid/plant complexes.