An artificial diet for continuous maintenance of Coccinella septempunctata adults (Coleoptera: Coccinellidae)

An artificial diet was developed for continuous maintenance of ladybird beetle (Coccinella septempunctata) adults. This beetle is an important biocontrol agent for greenhouse crops. Several biological parameters were measured and compared between colonies fed the artificial diet over eight successive generations, and colonies fed aphids. Weight, survival rates from 1st to 4th instar, pupation rate, and emergence rate were similar for both colonies, but the sex ratio (♀:♂) of the artificial diet-fed colony was higher than that of the aphid-fed colony. These results suggest that this artificial diet could meet the basic needs for the successive rearing of C. septempunctata adults. The prepared diet, which is semi-solid, provides suitable moisture for C. septempunctata, and the diet ingredients are inexpensive and relatively easy to prepare. This diet and rearing technique offers a very useful approach for mass production of C. septempunctata.     

The influence of some honeybee products as a diet substitute on the different stages of Coccinella undecimpunctata L. in Egypt

Several natural diets of honeybee products were tested for rearing Coccinella undecimpunctata L. under laboratory conditions. Two groups of tested adult and larval insect have been fed on the tested diets, the first group were fed on the tested honey bee products diets without adding aphids, while the second group fed on the same honeybee products with aphids added. The oviposition period in the first group was significantly shorter than control. The tested diets supplied with royal jelly + aphid and/or pollen grains + aphid caused the longest oviposition period. Diets of aphid alone and bee honey with aphid recorded the shortest oviposition period. Fecundity of ladybird was the highest in the tested diets with aphids than diets without aphid and/or aphid diet alone. The durations of larval and pupal stages were significantly different when compared with the diets with or without aphid and/or aphid diet alone. Further results indicated that the diets of royal jelly caused the shortest duration in larval and pupal stages.     

Ladybird population dynamics in potato: comparison of native species with an invasive species, Harmonia axyridis

Following the detection of the harlequin ladybird, Harmonia axyridis, in 2003 in potato crops in Belgium, a study was carried out between 2004 and 2006 on the phenology of this species compared to native species in potato. The results confirmed the success of H. axyridis, with high population levels in 2004 and 2005. In 2006, aphid populations were very low and no H. axyridis larvae were sampled in potato, but the indigenous species Coccinella septempunctata and Propylea quatuordecimpunctata were detected. A species by species comparison of the date of first larvae detection, the larvae population peak, and the difference between this peak and the aphid population peak was performed. Results showed a clear correlation between C. septempunctata and P. quatuordecimpunctata and potato aphids, with a delay of 3.5 and 6.5 days between the aphid and ladybird population peaks for the two native species. H. axyridis arrived 7–8 days after the two indigenous species and the larval peak population occurred 15.8 days after the aphid population peak. This meant that H. axyridis had to complete its larval development with very low aphid populations or even with no aphids at all. The reason for its late arrival and the possible food resources used by H. axyridis larvae are discussed.     

Predation and avoidance behaviour in aphid‐ladybird interactions of native and invasive ladybirds in Europe

1. While detrimental effects of invasive predators on native species are well documented, we often lack a mechanistic understanding of the invasion success. Lack of prey avoidance behaviour can lead to higher consumption rates by invasive predators compared to native predators. This competitive advantage is expected to contribute to the invasion success of non‐native predators.
2. We compared aphid consumption and cue avoidance behaviour of aphids between four native ladybird species (Coccinella septempunctata, Adalia bipunctata, Propylea quatuordecimpunctata, and Hippodamia variegata) and the invasive Asian ladybird Harmonia axyridis.
3. The invasive H. axyridis and the native C. septempunctata consumed more aphids than the three smaller native ladybird species. In line with our expectations, aphids avoided leaves bearing cues of most native ladybird species but not of the invasive H. axyridis.
4. Our results indicate that body size rather than ladybird origin determined aphid predation rates. The lack of aphid avoidance behaviour towards cues of H. axyridis indicates that they were not able to recognise the chemical cues of the invasive predator.
5. Relatively large body size and the absence of cue avoidance in aphids might benefit the invasive H. axyridis, particularly in comparison to smaller native ladybird species. The absence of avoidance behaviour in aphids might lead to even higher predation rates of H. axyridis under more natural conditions.

     

Effects of Aphid Prey on Larval Development and Mortality of Adalia bipunctata and Coccinella septempunctata (Coleoptera: Coccinellidae)

Developmental time and mortality rate of Adalia bipunctata (L.) and Coccinella septempunctata L. (Coleoptera: Coccinellidae) were determined when feeding on five aphid species. Metopolophium dirhodum (Walker), Sitobion avenae (F.), Rhopalosiphum padi (L.), Hyalopterus pruni (Geoffr.) and Myzus cerasi F. (Homoptera: Aphididae) are widespread in Tekirda?, Turkey. Tests were carried out in a controlled environmental chamber (25±1°C temperature, 65±5% relative humidity and 16 h light:8 h dark period). Developmental times for A. bipunctata and C. septempunctata larvae varied significantly depending on species of aphid prey (P<0.05). Development time (±S.E.) varied from 17.50±0.84 to 20.83±1.60 days for C. septempunctata and 16.7±0.76 to 20.7±1.03 days for A. bipunctata. Mortality of A. bipunctata (50%) and C. septempunctata (63%) were highest on H. pruni.     

Influence of six aphid prey species on development and reproduction of a ladybird beetle, Coccinella septempunctata

Pre-imaginal development, immaturesurvival, and reproduction of a ladybird, Coccinella septempunctata Linnaeus, werestudied in response to six aphid species, Aphiscraccivora Koch, Aphis gossypii Glover,Aphis nerii Boyer de Fonscolombe,Lipaphis erysimi (Kaltenbach), Myzuspersicae (Sulzer) and Uroleuconcompositae (Theobald) to quantify theirrelative suitability as prey. Pre-adultdevelopment was shortest (13.93 ± 0.12 days)when fed on L. erysimi and longest(22.85 ± 0.10 days) on A. nerii. Immaturesurvival, adult emergence, growth index,relative growth rate, development rate, maleand female longevity, oviposition period,fecundity and hatching percent were maximal, i.e. 73.47 ± 0.89%, 90.07 ± 1.43%,8.62 ± 0.23, 1.52 ± 0.02, 0.07,81.10 ± 1.26 days, 85.70 ± 1.45 days,69.80 ± 1.32 days, 1764.10 ± 8.46,and 87.88 ± 1.05, respectively when C.septempunctata were fed on L. erysimi.The same parameters were minimal, i.e.43.86 ± 1.33%, 71.65 ± 2.75%,2.02 ± 0.08, 0.49 ± 0.02, 0.04,44.40 ± 1.39 days, 53.50 ± 1.00 days,16.40 ± 0.60 days, 203.20 ± 11.83, and48.68 ± 2.06, respectively on A. nerii. Theweights of different ladybird life stages weremaximal after feeding on L. erysimi and minimalon A. nerii. Regression analyses of thedata revealed linear relationships betweendevelopment rate and weight of adult; dailyprey consumption and relative growth rate; logweight of adult male and female; and longevityand fecundity of female. On the basis of thesefindings, the order of suitability of aphidspecies for C. septempunctata is L. erysimi >M. persicae > A. craccivora > A. gossypii >U. compositae > A. nerii. Thus, the presentinformation can be utilized for the massrearing of C. septempunctata by supplyingthe best food and can also help in theprediction of the relative abundance of theladybird on different aphid infestations in thefields.     

Effect of temperature on development, growth and feeding of Coccinella septempunctata and Hippodamia convergens reared on the tobacco aphid, Myzus persicae nicotianae

Preimaginal development, mortality, aphid consumption rate, and size and weight upon reaching the adult stage of the aphidophagous coccinellids Hippodamia convergens Guérin-Méneville and Coccinella septempunctata L. collected from Karditsa, central Greece, were examined at four constant temperatures (14, 17, 20 and 23 °C) and L16:D8. The coccinellids fed on the tobacco aphid, Myzus persicae nicotianae Blackman. Egg, larval and pupal mortality was highest at 14 °C reaching 85.0, 73.8 and 29.4% in H. convergens and 49.3, 75.4 and 58.8% in C. septempunctata, respectively. Total preimaginal development ranged from 57.2 to 70.4 days at 14 °C, and to 16.9 and 22.1 days at 23 °C in H. convergens and C. septempunctata, respectively. Heavier and larger adults of H. convergens were obtained at 17 and 20 ° C. In C. septempunctata temperature did not affect adult weight while the lowest size was observed at 14 and 17 °C. Day-degrees requirements for preimaginal development in H. corvengens were 212.9 above a developmental threshold of 11.0 °C. The corresponding values for C. septempunctata were 281.5 and 10.7 °C. In H. convergens total and daily aphid consumption ranged from 46.8 aphids at 14 °C to 85.0 aphids at 23 °C and from 1.5 aphids at 14 °C to 9.2 aphids at 23 °C, respectively. The corresponding values for C. septempunctata were 112.0 and 2.7 at 14 °C and 157.7 and 12.4 at 23 °C. The results show the high potential of both predators as biological control agents against the tobacco aphid. The knowledge obtained could be essential for their appropriate use and for the improvement of mass rearing systems.     

Toxicity of the aphid Aulacorthum magnoliae to the predator Harmonia axyridis (Coleoptera: Coccinellidae) and genetic variance in the assimilation of the toxic aphids in H. axyridis larvae

Larvae of the ladybird Harmonia axyridis are reported to suffer high mortality when they are fed with the aphid Aulacorthum magnoliae associated with the elder Sambucus sieboldiana. In the present study we first examined whether aphid toxicity to H. axyridis was altered when the aphids were reared on different host plants, and second whether some ladybird families could adapt specially to the toxic aphids. Ladybird larvae from each egg batch were divided into two groups; one group was fed with A. magnoliae reared on potato, and the other group was fed with A. magnoliae reared on elder. The ladybird larvae fed with elder aphids suffered significantly higher mortality and grew more slowly than did larvae fed with potato aphids. This result indicates that A. magnoliae aphids absorbed toxic substances or their precursors from S. sieboldiana. We suggest that host association of A. magnoliae with the primary host plant S. sieboldiana has been maintained because of the advantage of deterring predation. Significant and positive correlation was detected across H. axyridis sib families between the mean weight of larvae fed with elder aphids and the mean weight of larvae fed with potato aphids. The hypothesis that some ladybird families adapt specially to the toxic aphids was not supported. However, the available evidence showed that a large amount of genetic variance in performance is maintained in a wild population of the ladybird H. axyridis.     

Negative dietary effects of Colorado potato beetle eggs for the larvae of native and introduced ladybird beetles

Aphidophagous ladybird beetles (Coleoptera: Coccinellidae) are attracted to and feed heavily on aphids, but many species will also feed opportunistically on other prey that they encounter. In potatoes (Solanum tuberosum L.) in Washington State, USA, coccinellids feed on both green peach aphids (“GPA,” Myzus persicae Sulzer) and eggs of the Colorado potato beetle (“CPB,” Leptinotarsa decemlineata Say). The guild of aphidophagous ladybirds includes two native species, Hippodamia convergens Guérin-Méneville and Coccinella transversoguttata Brown. Recently, an introduced species, Coccinella septempunctata L., has invaded and apparently displaced its native congener. A second exotic, Harmonia axyridis Pallas, has colonized the area and is becoming more abundant. We compared larval development of each species on a monotypic diet of GPA, a monotypic diet of CPB eggs, or a mixed diet of both GPA and CPB eggs. Our goal was to answer two questions: (1) do larvae of the four ladybird species benefit from including CPB eggs in their diet and (2) do the four ladybird species differ in their ability to utilize CPB eggs as prey? No larva of any species completed development on a pure diet of CPB eggs, and survivorship was highest for all species when they fed on a pure diet of GPA. One native species, H. convergens, and one exotic species, H. axyridis, exhibited significantly lower survivorship on a mixed diet of both CPB eggs and GPA, compared to a pure GPA diet; H. axyridis also took longer to develop from egg to adult when both prey were provided. Survivorship of the two Coccinella spp. was not altered by the inclusion of CPB eggs with GPA, although CPB eggs lengthened the development time of C. transversoguttata. Adult size was not consistently affected by diet for any of the coccinellids. Overall, no ladybird species benefited from the inclusion of potato beetle eggs in its diet. The two Coccinella species responded similarly to the inclusion of CPB eggs, and so we would not expect any difference in the success of coccinellid larval development in potato fields following the replacement of C. transversoguttata by C. septempunctata. Hippodamia convergens and H. axyridis, the two species whose survivorship was depressed by combining CPB egg and aphid prey, were also the two species that consumed the greatest number of CPB eggs during successful larval development. A comparison of total egg consumption by each species cohort suggested that displacement of the other species by H. axyridis would not alter CPB biological control, because the higher per capita feeding rate by H. axyridis larvae compensated for individuals’ greater mortality risk on a diet including CPB eggs.     

Preliminary observations on the population development of anystid and free-living mesostigmatic mites in a cereal field in Southern England

The effect of manipulating cereal aphid population development, by either adding aphids or spraying with an aphid-specific insecticide, on the predatory mite fauna was investigated. Populations of the anystid mite, Anystis baccarum (L.) were very small until late June when numbers built up rapidly at the time of the aphid population decline. There was however, no significant relationship between aphids and numbers of mites over the whole season. Preliminary electrophoretic analysis of two enzymes in the gut of anystid mites indicated that they may have been feeding on thrips, Limothrips cerealium Haliday. The free-living soil Mesostigmata had two population peaks, each related, the first positively and the second negatively to aphid populations.     

Essential versus alternative foods of insect predators: benefits of a mixed diet

Although many predatory insects appear to be opportunistic generalists in their selection of prey, only a subset of prey species may in fact serve as “essential foods” capable of supporting immature growth and adult reproduction. It has been suggested that other, “alternative foods” serve only to maintain the predator when essential foods are not available, but little research has evaluated the significance of a mixed diet of essential and alternative foods for predator growth or reproduction. Here we test the general hypothesis that although alternative prey may be inadequate to support reproduction when consumed alone by adult predators, consumption of such prey may enhance the predator's reproductive output when the predator also has access to essential prey. We compared egg production by two aphidophagous lady beetles, Coccinella septempunctata and C. transversoguttata, provided with diets of aphids (essential prey) and weevils (alternative prey). As predicted, female predators produced greater numbers of eggs when a diet of pea aphids in limited number was supplemented by alfalfa weevil larvae. The predators laid no eggs when provided only with weevils or only with sugar. But once aphids were added to the diet, females of C. transversoguttata (but not C. septempunctata) laid eggs in greater numbers when they had fed previously on weevils than on sugar. Females of both species also produced eggs in modest numbers when provided with both weevils in excess and sugar, but this diet supported a lower rate of egg production than did a diet of weevils in excess plus a limited number of aphids. Although C. septempunctata has a longer history of association with the alfalfa weevil than does C. transversoguttata, the former species was not more effective in exploiting this alternative prey in support of reproduction. The tendency of generalist predators such as adult lady beetles to consume alternative as well as essential prey probably enhances considerably their ability to capitalize on short-lived and scattered opportunities as they seek out suitable sites in which to reproduce. Received: 11 May 1998 / Accepted: 1 May 1999     

Bottom–up regulates top–down: the effects of hybridization of grass endophytes on an aphid herbivore and its generalist predator

The ecological consequences of hybridization of microbial symbionts are largely unknown. We tested the hypothesis that hybridization of microbial symbionts of plants can negatively affect performance of herbivores and their natural enemies. In addition, we studied the effects of hybridization of these symbionts on feeding preference of herbivores and their natural enemies. We used Arizona fescue as the host‐plant, Neotyphodium endophytes as symbionts, the bird cherry–oat aphid as the herbivore and the pink spotted ladybird beetle as the predator in controlled experiments. Neither endophyte infection (infected or not infected) nor hybrid status (hybrid and non‐hybrid infection) affected aphid reproduction, proportion of winged forms in the aphid populations, aphid host‐plant preference or body mass of the ladybirds. However, development of ladybird larvae was delayed when fed with aphids grown on hybrid (H+) endophyte infected grasses compared to larvae fed with aphids from non‐hybrid (NH+) infected grasses, non‐hybrid, endophyte‐removed grasses (NH?) and hybrid, endophyte‐removed (H?) grasses. Furthermore, adult beetles were more likely to choose all other types of grasses harboring aphids rather than H+ infected grasses. In addition, development of ladybirds was delayed when fed with aphids from naturally uninfected (E?) grasses compared to ladybirds that were fed with aphids from NH+ and NH? grasses. Our results suggest that hybridization of microbial symbionts may negatively affect generalist predators such as the pink spotted ladybird and protect herbivores like the bird cherry–oat aphids from predation even though the direct effects on herbivores are not evident.     

Interspecific interactions in a tri-trophic arthropod system: effects of a spider on the survival of larvae of three predatory ladybirds in relation to aphids

The effects of the crab spider, Misumenops tricuspidatus (Fabricius), on the larval survival of three ladybird species, Harmonia axyridis Pallas, Coccinella septempunctata L., and Propylea japonica L., in relation to aphids were investigated in the laboratory. Predation by the spider on the three ladybird species differed. All the larvae of C. septempunctata, none of H. axyridis, and an intermediate number of P. japonica were attacked and eaten by the spider. All the larvae of H. axyridis suffered mortality due to cannibalism or starvation in the treatments with and without a spider. In case of C. septempunctata, however, mortality in the early instars was significantly greater in the treatment with a spider than without a spider and no larvae developed into pupae due to predation. In the treatment without a spider, the majority of the larvae in the former treatment suffered mortality due to cannibalism or starvation, and only 13.3% of larvae developed into the adult stage. In the case of P. japonica, mortality was mainly attributed to predation in the treatment with a spider and only 26.7% became adult. In comparison, 86.7% of larvae survived to the adult stage in the treatment without a spider. In addition, in both H. axyridis and C. septempunctata, the development of young larvae was significantly slower in the presence of a spider, but this was not the case with the older larvae of H. axyridis, which indicates that the effect of the spider on larval development changed with the developmental stage of the larvae in this species. However, the spider had no significant effect on the developmental time of P. japonica larvae. Although both the spider and the ladybirds significantly affected the number of aphids, they did not have an additive effect on aphid abundance. The interactions between the spider and the ladybirds, such as intraguild predation or competition, caused them to reduce aphid population density less than the ladybirds did on their own. The effect of the spider on the larval performance of three predatory ladybirds was found to be unequal in terms of their vulnerability to predation and rate of larval development and it depended on the species and developmental stage of the ladybird.     

Infection with an insect virus affects olfactory behaviour and interactions with host plant and natural enemies in an aphid

Aphid ecology and population dynamics are affected by a series of factors including behavioural responses to ecologically relevant chemical cues, capacity for population growth, and interactions with host plants and natural enemies. Using the aphid Rhopalosiphum padi (L.) (Homoptera: Aphididae), we showed that these factors were affected by infection with Rhopalosiphum padi virus (RhPV). Uninfected aphids were attracted to odour of uninfected aphids on the host plant, an aggregation mechanism. However, infected aphids were not attracted, and neither infected nor uninfected aphids were attracted to infected aphids on the plant. Infected aphids did not respond to methyl salicylate, a cue denoting host suitability. Infected aphids were more behaviourally sensitive to aphid alarm pheromone, and left the host plant more readily in response to it. RhPV reduced the lifespan and population growth rate of the aphid. The predacious ladybird, Coccinella septempunctata (L.) (Coleoptera: Coccinellidae), consumed more infected aphids than uninfected aphids in a 24‐h period, and the aphid parasitoid Aphidius ervi Haliday (Hymenoptera: Aphidiidae) attacked more infected than uninfected aphids. However, the proportion of mummies formed was lower with infected aphids. The results represent further evidence that associated organisms can affect the behaviour and ecology of their aphid hosts.     

Suitability of greenbugs, cotton aphids, andHeliothis virescens eggs for development and reproduction ofOrius insidiosus

Development, fecundity, and longevity of the predator,Orius insidiosus (Say) (Heteroptera: Anthocoridae), when reared on greenbug [Schizaphis graminum (Rondani)], cotton aphid (Aphis gossypii Glover), or eggs of tobacco budworm [Heliothis virescens (F.)] with green beans or water were examined. Developmental time was shortest when predators were reared onH. virescens eggs and beans and longest when reared on cotton aphids and water. Predators were most fecund when fedH. virescens eggs. The inclusion of beans in the nymphal diet further enhanced fecundity when fed eggs. Longevity of both females and males was significantly shorter when reared on aphids than on eggs. Beans in the nymphal diet enhanced longevity of female predators only in combination with budworm eggs. FemaleO. insidiosus were largest when reared onH. virescens eggs. Addition of green beans in aphid treatments resulted in increased size ofO. insidiosus when compared to aphids and free water.     

INFLUENCE OF TEMPERATURE ON CERTAIN BIOLOGICAL ATTRIBUTES OF A LADYBEETLE COCCINELLA SEPTEMPUNCTATA LINNAEUS

Influence of temperature on certain biological attributes of an aphidophagous ladybeetle, Coccinella septempunctata Linnaeus, feeding on mustard aphid, Lipaphis erysimi (Kaltenbach), at five different temperatures, viz. 20, 25, 27, 30 and 35°C was investigated. Its developmental period was shortest (11.7 ± 0.09 days) at 35°C and longest (20.6 > 0.35 days) at 20°C. Developmental rate increased with increase in temperature. Hatching percent, larval survival, adult emergence and growth index were maximum at 30°C and minimum at 20°C. Oviposition period and fecundity were highest at 30°C and lowest at 20°C. A positive linear relationship exists between temperature and developmental rate and negative correlation between the duration of immature life stages and temperature. The proportion of developmental period allocated to each immature stage was found to be similar at each temperature regime. Thus, 30°C was found as the most suitable for C. septempunctata amongst the five temperatures tested.     

Biology ofAllothrombium pulvinum (Acariformes: Trombidiidae), a potential biological control agent of aphids in China

Allothrombium pulvinum Ewing is a common natural enemy of aphids and other small arthropods in China. It is univoltine species. The eggs hatch in spring, nymphs emerge in early summer, and adults emerge in autumn. Larvae are ectoparasites of aphids, whereas deutonymphs and adults are free-living predators of aphids, spider mites and some lepidopterous eggs. Adults hibernate in soil during winter and females lay their eggs in spring. In the laboratory the duration of immature stages is 74 days at 20–30°C. Larvae kill their host within 1–3 days when the mite load of the host is 2 or more. They decrease the reproductive rate of adult aphids and arrest the development of early-instar aphid nymphs when the mite load is 1. Larvae are capable of controlling the host population growth when the level of parasitism is high. Owing to its wide occurrence,A. pulvinum could be a potential candidate in the biological control of aphids. It is suggested that its impact on the host pest be evaluated in IPM programs in the future.     

Weak responses to dietary enrichment in a specialized aphid predator

Some ladybeetles are specialist predators of aphids, coccids or other prey, although they often eat a variety of species from their focal prey taxon. In addition, the diet is often supplemented with alternative prey. How larvae of the aphidophagous Coccinella septempunctata L. utilize a non‐aphid alternative prey (fruit‐fly larvae Drosophila melonogaster Meigen) is compared with adequate (i.e. high‐quality) aphid prey provided alone (monotypic diet) or in mixed diets. The alternative prey are presented either nutrient‐enriched (i.e. raised on dog food supplemented medium) or not (raised on pure medium). Ladybird performance (survival, growth and development) is poor on the pure fly larvae diets, and also reduced when given mixed diets compared with the pure aphid diet. Nutrient enrichment of the fly larvae has no positive effects. The physiological background for the differences in food value, as indicated by performance in life‐history parameters, is a strong pre‐ingestive effect (i.e. reduced consumption of fly larvae compared with aphids) and a post‐ingestive effect (i.e. reduced utilization of assimilated larval fly tissue), whereas the assimilation efficiency of the consumed fly larvae is as high as that of aphids. The results show a physiological trade‐off resulting from prey specialization that reduces the possibility of utilizing alternative prey when the availability of aphids is scarce. Connected with this is a high robustness against variation in prey nutrient diversity and composition; the ladybird shows little positive response to dietary mixing (i.e. neither mixing of adequate aphids, nor of aphids and alternative prey) or to nutrient enrichment of prey. This contrasts with the results from generalist predators (spiders), where similar treatments lead to strong effects on life‐history parameters.     

Intraguild predation and successful invasion by introduced ladybird beetles

Introductions of two ladybird beetle (Coleoptera: Coccinellidae) species, Coccinella septempunctata and Harmonia axyridis, into North America for aphid biocontrol have been followed by declines in native species. We examined intraguild predation (IGP) between larvae of these two exotic species and larvae of the two most abundant native coccinellids in eastern Washington State, C. transversoguttata and Hippodamia convergens. In pairings between the two native species in laboratory microcosms containing pea (Pisum sativum) plants, neither native had a clear advantage over the other in IGP. When the natives were paired with either Harmonia axyridis or C. septempunctata, the natives were more frequently the victims than perpetrators of IGP. In contrast, in pairings between the exotic species, neither had an IGP advantage, although overall rates of IGP between these two species were very high. Adding alternative prey (aphids) to microcosms did not alter the frequency and patterns of relative IGP among the coccinellid species. In observations of encounters between larvae, the introduced H. axyridis frequently survived multiple encounters with the native C. transversoguttata, whereas the native rarely survived a single encounter with H. axyridis. Our results suggest that larvae of the native species face increased IGP following invasion by C. septempunctata and H. axyridis, which may be contributing to the speed with which these exotic ladybird beetles displace the natives following invasion.     

Rhythm of honeydew excretion by the tea aphid and its attraction to various natural enemies

The rhythm of honeydew excretion by the tea aphid Toxoptera aurantii (Boyer) and its attraction to following 9 species (or subspecies) of beneficial insects, Aphidius sp., Chrysopa sinica Tjeder, Chrysopa septempunctata Wesmael, Sphaerophoria menthastri L., Coccinella septempunctata L., Leis axyridis (Pallas) ab. bimaculata Hemmelmann, L. axyridis (Pallas) ab. conspicua Faldermaenn, L. axyridis (Pallas) var. spectabilis Faldermaenn and L. axyridis (Pallas) var. novemdecimpunctata Faldermaenn, were investigated. Forty-five wingless virginoparae nymphs, reproduced by the same wingless virginogenia adult within 1 h, were introduced onto tea seedlings with one aphid per seedling. The honeydews excreted from different instars of nymphs and adults were collected under 21°C, 85% RH, 3500 lx and 12 L:12 D photoperiod. It took 32.4 d ±5.8 d for the tested tea aphids to complete the development of their nymph and adult stages, during which 325.6±35.8 droplets (ca. 41.98 μl ±6.14 μl and 45.34 mg ± 8.76 mg) of honeydews were secreted. During the 1st and 4th instars, there was a logistic regression relationship between the amount of honeydews excreted and the time (days). The honeydew secretion during the first two instars was less than that from the later instars. Adult aphids survived for 22.0 d ± 0.0 d, and excreted 176.31 ± 22.38 droplets (ca. 30.38 μl ± 5.32 μl) of honeydews in a rate of 1 drop per ca. 30–50 min for 5–8 h with a pause for 2–5 h before next secretion series. A batch of forty-five virginoparae female adults, reproduced by the same virginoparae female adult within 1 h, were introduced onto the tea seedlings (one aphid/seedling) under 13–21°, 85% RH, 3500 lx and 12 L:12 D photoperiod. Temperature showed a significant effect on the amounts of honeydews excreted within the range of 13–21°. Honeydews excreted by the aphids significantly increased the searching and retention time of the tested 9 species of natural enemies in a positive dose-response fashion. The searching times of Aphidius sp. and S. menthastri were the longest and the shortest, respectively, among all the 9 species, while the searching and retention time of L. axyridis (Pallas) var. spectabilis was the longest among the four varieties of L. axyridis. Tea aphid oneydew is considered as an important contact kairomone for the tested natural enemies.