Differential sensitivity of an invasive and an indigenous ladybeetle to two reduced‐risk insecticides

The invasive multicoloured Asian lady beetle, Harmonia axyridis (Pallas), and the indigenous twelve spotted lady beetle, Coleomegilla maculata DeGeer (Col., Coccinellidae), are two important generalist predators commonly found in apple orchards in Quebec, Canada. Both species are exposed to two reduced‐risk insecticides, recently adopted to control codling moth, Cydia pomonella (L.) (Lep., Tortricidae) in south‐eastern Canada. Chlorantraniliprole (Altacor^® 35 WG), an anthranilic diamide insecticide, causes paralyses of the muscle cells by interfering with the insect ryanodine receptors, whereas novaluron (Rimon^® EC 10), a benzoylphenyl urea, inhibits the chitin synthesis. The aim of this study was to compare the sensitivity of both the invasive ladybeetle H. axyridis and the indigenous C. maculata to reduced‐risk insecticides through the assessment of lethal effect on eggs and larvae following topical contact, ingestion of treated prey and exposure to fresh residues, at field rates (50.75 g a.i./ha chlorantraniliprole and 100 g a.i./ha novaluron) in laboratory conditions. Eggs of both species were not affected. Following 6 days of residual contact, chlorantraniliprole and novaluron caused more than 98% mortality to larvae of both ladybeetle species. In topical contact and ingestion trials, chlorantraniliprole caused less than 18% mortality to larvae of the two species after 6 days following exposure. Novaluron had a drastically different impact on the two predators. It did not affect the indigenous C. maculata, whereas it killed 91% and 96% of H. axyridis individuals after 6 days, respectively, following topical contact and ingestion. These results illustrate a differential sensitivity to novaluron between two relatively close species (subfamily Coccinellinae), a potential impact on the invasion process by H. axyridis, and consequently on the ladybeetle assemblage in the field.     

A one‐step,single tube,duplex PCR to detect predation by native predators on invasive Bemisia tabaci MEAM1 and Frankliniella occidentalis

Bemisia tabaci (Gennadius) MEAM1 (Hemiptera: Aleyrodidae) and Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), two important invasive species, are serious agricultural pests. In this study, a one‐step, single tube, duplex polymerase chain reaction (PCR) procedure was developed to allow rapid, specific, and sensitive identification of B. tabaci MEAM1 and F. occidentalis in predator guts. The system and conditions used for the duplex PCR were optimized. The species specificity of the duplex PCR determined by comparison against non‐targets that might interact with B. tabaci MEAM1 and F. occidentalis showed that oligonucleotide primers amplified nuclear gene target sequences present only in B. tabaci MEAM1 or F. occidentalis. The limits of detection were 9.53 ng μl^?1 for B. tabaci MEAM1 and 8.94 ng μl^?1 for F. occidentalis. Within a field cage study, in which predators Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) and Orius sauteri (Poppius) (Hemiptera: Anthocoridae) were allowed to feed on B. tabaci MEAM1 and F. occidentalis for 10 h, the B. tabaci MEAM1 DNA was detectable in 100% of H. axyridis and O. sauteri, and F. occidentalis DNA was detectable in 80% of H. axyridis and 90% of O. sauteri; this implicated that B. tabaci MEAM1 and F. occidentalis remains could be detected in native predator guts simultaneously. The accuracy and reliability of the assay suggested strongly that the duplex PCR optimized for B. tabaci MEAM1 and F. occidentalis is sensitive and specific for both invasive insects and is therefore useful in early diagnosis and monitoring of B. tabaci MEAM1 and F. occidentalis infections, and can be used to identify domestic predator species and food web relationships.     

Weak photoperiodic response facilitates the biological invasion of the harlequin ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae)

Photoperiodic regulation of reproductive diapause in two invasive and two native populations of Harmonia axyridis and in one native population of Harmonia yedoensis was investigated in laboratory at 20°C, five photoperiods (day length of 10, 12, 14, 16 and 18 h) and two diets: (i) eggs of the Angoumois grain moth Sitotroga cerealella and (ii) the green peach aphid, Myzus persicae. Laboratory strains originated from native populations of H. axyridis from Irkutsk province of Siberia and H. yedoensis from South Korea showed a strong photoperiodic response: under short photoperiods (10–14 h and 10–12 h for H. axyridis and H. yedoensis, correspondingly), all females which fed on eggs and most of those fed on aphids did not start to lay eggs during 40 days after emergence, while under long photoperiods, all females fed on aphids and most of those fed on eggs oviposited. The photoperiodic response of H. axyridis from South Korea was less strong: on the both diets, the range of the photoperiodic response (the difference in the proportion of ovipositing females between the treatments with long and short days) was ca 40%. In the European (Czech Republic) and in the Caucasian (Sochi region, Russia) invasive populations of H. axyridis, the photoperiodic response was very weak: the proportion of females that started oviposition (when fed on aphids) or at least reproductive maturation (when fed on eggs) during 40 days after emergence was close to 100%, independently of the photoperiodic conditions. Obviously, instead of a rapid micro‐evolutionary adaptation of the critical day length to a new climate, the invasive populations of the harlequin ladybird decrease their dependence on photoperiod and thus the weak photoperiodic response of SE Asian population of H. axyridis can be considered as a pre‐adaptation further developed during the invasion.     

Influence of prey quality on the fitness of two phenotypes of Harmonia axyridis adults

The performance of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) adults of the aulica and nigra phenotypes fed on Aphis fabae Scopoli and Myzus persicae (Sulzer) was compared by measuring their voracity, daily biomass consumption, daily weight gain, efficiency of food utilisation, and reproductive capacity. Our results demonstrated differences in the suitability of A. fabae and M. persicae for the two phenotypes of the predator. This suggests that either differences occur in the nutritive requirements of the predators, or in the nutritive value of the two prey species. Both A. fabae and M. persicae supported the growth and oviposition of the aulica and nigra phenotypes. Although nigra females consumed fewer M. persicae, they achieved the same daily weight gain as aulica females. The predator phenotypes consumed the same amount of A. fabae, but the daily weight gain of aulica females was higher than that of nigra. The two predator phenotypes had the same feeding efficiency when consuming M. persicae or A. fabae. The reproductive capacity of nigra females was higher, when this phenotype consumed A. fabae rather than M. persicae.     

Sublethal effect of beta‐cypermethrin on development and fertility of the Asian multicoloured ladybird beetle Harmonia axyridis

The Asian multicoloured ladybird beetle, Harmonia axyridis, is utilized as a major natural enemy of aphids in the field, greenhouses and orchards. However, it has been looked as invasive predator distributing in worldwide. To refine integrated pest management (IPM) against aphids, it is important to evaluate the effects of insecticides on physiology and behaviour of the high adapted predators. Beta‐cypermethrin, a broad‐spectrum insecticide, not only kills aphids at lethal concentrations but also affects natural enemy of aphids. In our study, the age‐stage, two‐sex life table was used to evaluating sublethal effects of beta‐cypermethrin on the predatory ladybird beetle H. axyridis. In the parent generation, the pre‐oviposition period of H. axyridis was significantly shortened (8.93 days) after exposure to LC₅ beta‐cypermethrin (5% lethal concentration) as compared with control (10.06 days). However, the oviposition period was significantly longer (46.17 days instead of 43.90 days), and fecundity (eggs per female) was significantly increased by 49.64% when compared with control. In the F₁ generation, the length of the juvenile stage was not affected, but the oviposition period increased significantly (38.19 days compared to 31.39 days in the control). This positive effect was translated to the fecundity that increased significantly by 62.27% as compared with control. According to the life‐table analysis, the intrinsic rate of increase (r_m) was significantly higher in treatment (0.140 per day) than that in the control (0.123 per day). In addition, the net reproductive rate (R₀) increased significantly by 91.53%. These results would be useful in assessing the overall effects of beta‐cypermethrin on H. axyridis and even for discussing the ecological mechanism of the unexpected extension of H. axyridis during IPM programme.     

Can chilling tolerance of C4 photosynthesis in Miscanthus be transferred to sugarcane?

The goal of this study was to investigate whether chilling tolerance of C₄ photosynthesis in Miscanthus can be transferred to sugarcane by hybridization. Net leaf CO₂ uptake (A_sat) and the maximum operating efficiency of photosystem II (Ф_PSII) were measured in warm conditions (25 °C/20 °C), and then during and following a chilling treatment of 10 °C/5 °C for 11 day in controlled environment chambers. Two of three hybrids (miscanes), ‘US 84‐1058’ and ‘US 87‐1019’, did not differ significantly from the chilling tolerant M. ×giganteus ‘Illinois’ (Mxg), for A_sat, and Φ_PSII measured during chilling. For Mxg grown at 10 °C/5 °C for 11 days, A_sat was 4.4 μmol m^?2 s^?1, while for miscane ‘US 84‐1058’ and ‘US 87‐1019’, A_sat was 5.7 and 3.5 μmol m^?2 s^?1, respectively. Miscanes ‘US 84‐1058’ and ‘US 87‐1019’ and Mxg had significantly higher rates of A_sat during chilling than three tested sugarcanes. A third miscane showed lower rates than Mxg during chilling, but recovered to higher rates than sugarcane upon return to warm conditions. Chilling tolerance of ‘US 84‐1058’ was further confirmed under autumn field conditions in southern Illinois. The selected chilling tolerant miscanes have particular value for biomass feedstock and biofuel production and at the same time they can be a starting point for extending sugarcane's range to colder climates.     

Does nitrogen limitation promote intraguild predation in an aphidophagous ladybird?

Reciprocal intraguild predation occurs between the two aphidophagous ladybird beetles Coccinella septempunctata L. and Harmonia axyridis Pallas (Coleoptera: Coccinellidae). However, its direction is asymmetrical; H. axyridis generally acts as an intraguild predator, and C. septempunctata as an intraguild prey. According to Denno and Fagan's prediction that nitrogen shortages in predators may promote intraguild predation, it was hypothesized that growth of intraguild predator H. axyridis is more limited by nitrogen than that of intraguild prey C. septempunctata, and that H. axyridis growth is enhanced by feeding on C. septempunctata compared to feeding on aphids. To determine nitrogen‐limited growth in H. axyridis, the following two predictions were examined. First, it was predicted that the nitrogen content of H. axyridis would be higher than that of C. septempunctata when both feed on aphids. However, nitrogen content did not differ between the two ladybirds. Second, it was predicted that nitrogen‐use efficiency of H. axyridis would be lower than that of C. septempunctata. However, there was no significant difference between species. These results did not support the hypothesis that growth of H. axyridis is more limited by nitrogen than that of C. septempunctata. In addition, the present study showed that dry mass and nitrogen growth of H. axyridis were not enhanced, but rather decreased, by eating high‐nitrogen C. septempunctata, compared to eating low‐nitrogen aphids. Overall, the present study did not support the hypothesis that nitrogen shortages in predators may promote intraguild predation.     

Coccinellid interactions mediated by vegetation heterogeneity

Environmental heterogeneity can have profound effects on agroecosystem function and it is important for improving ecosystem services such as biological control. Promoting system diversity via non‐crop plants is one method for increasing habitat heterogeneity within farmscapes. Non‐crop plants provide access to refuges and alternative food resources provide multiple benefits to enhance populations of arthropod predators. In this study, we examined the effects of small‐scale spatial structure on life‐stage specific interactions between the native coccinellid, Hippodamia convergensGuérin‐Méneville, and the exotic Harmonia axyridis (Pallas) (both Coleoptera: Coccinellidae), which overlap in spatial distribution in many crop systems. Squash [Cucurbita pepo L. (Cucurbitaceae)] and non‐crop mugwort [Artemisia vulgaris L. (Asteraceae)] plants with and without aphids were used as a model of spatial heterogeneity in micro‐ and mesocosm experiments. In response to factorial treatment combinations, we evaluated oviposition behavior, egg predation, larval survival, and larval predator‐prey and predator‐predator interactions. Adult H. convergens displayed higher foraging activity on aphids when exposed to complex habitats containing a non‐crop plant. In the presence of the exotic coccinellid, H. convergens preferred to deposit eggs on the non‐crop plant. Furthermore, a combination of spatial heterogeneity and prey availability reduced larval intraguild predation and cannibalism, and improved reproductive output of H. convergens by reducing intra‐ and interspecific egg predation. Our results provide evidence that life‐stage‐specific intraguild interactions are mediated by access to non‐crop plants. Thus, the introduction or maintenance of non‐crop plants has the potential to enhance coexistence of multiple natural enemies and improve top‐down control of pests.     

Hydrosols evaluation in pest control: insecticidal and settling inhibition potential against Myzus persicae (Sulzer)

The hydrosols are by‐products derived during the extraction of essential oils. Although essential oils have been widely evaluated for their insecticidal activities, the possible use of hydrosols in pest control has been almost unknown. The effects of the hydrosols of Origanum majorana (marjoram), Mentha pulegium (pennyroyal), and Melissa officinalis (lemon balm) on the survival and settling behaviour of the aphid pest Myzus persicae were investigated. The hydrosols were isolated using Clevenger hydrodistillation (i) with conventional heating (HD) and (ii) assisted by microwaves (MWHD). GC‐MS analysis showed that the volatiles occurring in the hydrosols were similar between the two techniques. Hydrosols were assayed for possible settling inhibitory effects on M. persicae in Petri dishes (15 cm diameter). In each dish, a sprayed together with an unsprayed eggplant leaf piece was placed. An adult aphid <24 h old was released on the treated leaf and its path length was recorded during the initial 10 min. Then, its position (on the treated or untreated leaf) was recorded 10, 20, 30, 60 min and 24 h after spraying. M. officinalis HD hydrosol resulted in fourfold increase of the path length compared with the control (deionized water). The last observation (24 h) revealed that M. officinalis and M. pulegium HD hydrosols had the strongest inhibitory effect. Additionally, O. majorana hydrosols caused 10–15% aphid mortality after 24 h. In this study, the wider use of the MWHD technique is further supported as in addition to its well‐known advantages (i.e. shorter distillation time, less energy consumption), the effects of the hydrosols produced are comparable to those obtained by HD. Most importantly, the results clearly showed that the potential of hydrosols in pest control ought not to be ignored and should attract the interest of future studies.     

Pea aphid dropping behavior diminishes foraging efficiency of a predatory ladybeetle

Antipredator defensive behaviors are a well‐studied and often crucial part of prey life histories, but little has been done to quantify how such behaviors affect natural enemies, their foraging, and their effectiveness as biological control agents. We explored how the generalist predatory coccinellid Harmonia axyridis Pallas (Coleoptera: Coccinellidae) affects the dropping behavior of the pea aphid, Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), and in turn, how that defensive behavior affects the foraging efficiency of the predator. Experimental arenas that allowed or prevented pea aphid dropping were compared to determine how dropping influences the foraging of multiple life stages of H. axyridis: second instars, fourth instars, and adults. Dropping reduced predation on aphids by all ladybeetle life stages. Despite older predators inducing more dropping, aphid dropping reduced predation by approximately 40% across all ladybeetle life stages. Aphid dropping and predator consumption of aphids were both correlated with how much the predator moved, which also increased with predator life stage. We suggest that the high rates of dropping induced by H. axyridis and the subsequent decrease in H. axyridis foraging efficiency may partially explain why H. axyridis is less effective at controlling pea aphids than it is at controlling other aphid species that do not drop.     

Effects of UVB Radiation on competition between the bloom‐forming cyanobacterium Microcystis aeruginosa and the Chlorophyceae Chlamydomonas microsphaera1

The growth, photosynthetic characteristics, and competitive ability of three algal strains were investigated under different doses of ultraviolet‐B (UVB) radiation (0, 0.285, and 0.372 W · m^?2). The organisms were the toxic bloom‐forming cyanobacterium Microcystis aeruginosa FACHB 912, nontoxic M. aeruginosa FACHB 469, and the green microalga Chlamydomonas microsphaera FACHB 52. In monocultures, the growth of all three strains was inhibited by UVB. In mixed cultures, enhanced UVB radiation resulted in decreased percentages of the two M. aeruginosa strains (19%–22% decrease on d 12 of the competition experiment). UVB radiation resulted in increased contents of chlorophyll a, b, and carotenoids (CAR) in C. microsphaera, and decreased contents of allophycocyanin (APC) or phycocyanin in the two Microcystis strains. All three strains showed increased levels of UVabsorbing compounds and intracellular reactive oxygen species under 0.372 W · m^?2 UVB radiation, and decreased light compensation points, dark respiratory rates, and maximal quantum efficiency of PSII. After a 20 h recovery, the photosynthetic oxygen evolution of C. microsphaera was restored to its maximum value, but that of Microcystis strains continued to decrease. Nonphotochemical quenching was increased by UVB radiation in C. microsphaera, but was unaffected in the two M. aeruginosa strains. Our results indicated that C. microsphaera has a competitive advantage relative to Microcystis during exposure to UVB irradiation.     

‘Myrosin cells’ are not a prerequisite for aphid feeding on oilseed rape (Brassica napus) but affect host plant preferences

The enzyme myrosinase (EC 3.2.3.1.147) is present in specialised myrosin cells and forms part of the glucosinolate–myrosinase system, also known as ‘the mustard oil bomb’, which has an important role in the defence system of cruciferous plants against insect pests. Transgenic Brassica napus MINELESS have been produced by transgenic ablation of myrosin cells. This prompted us to investigate the importance of myrosin cells in plant–aphid interactions. In order to study this, we challenged transgenic MINELESS and wild‐type cultivar Westar seedlings with the aphids Brevicoryne brassicae (a specialist) and Myzus persicae (a generalist). Our study included aphid free‐choice and aphid fecundity experiments. Data from these experiments showed that B. brassicae prefers wild‐type seedlings and M. persicae prefers MINELESS. B. brassicae and M. persicae showed significant variation in establishment on plants regardless of whether they were wild type or MINELESS and also differed significantly in affecting plant parts. Myrosinase activity in MINELESS control seedlings was 83.6% lower than the wild‐type control seedlings. Infestation with either of the two aphid species induced myrosinase levels in both wild‐type and MINELESS seedlings. Infestation with M. persicae reduced the concentration of most glucosinolates while B. brassicae had the opposite effect. B. brassicae enhanced the formation of glucosinolate hydrolysis products both in wild‐type and MINELESS seedlings. However, M. persicae decreased All ITC but increased 3,4ETBut NIT in wild‐type seedlings. Taken together, the investigation shows that the presence of myrosin cells affects the preference of generalist and specialist aphid species for Brassica napus plants.     

Variation in Aggressiveness Components in the Hemileia vastatrix Population in Brazil

The Pucciniomycete fungus Hemileia vastatrix causes leaf rust on coffee trees. The pathogen is responsible for considerable yield losses in susceptible coffee cultivars if appropriate management strategies are not implemented. Rapid spread and epidemics of rust fungi are usually associated with the emergence of new races of the pathogen that overcome resistance or with the emergence of more aggressive populations of the pathogen. In Brazil, coffee production is dominated by susceptible cultivars of Coffea arabica and Coffea canephora. We assessed aggressiveness in 46 populations of H. vastatrix from Minas Gerais and Espírito Santo, two of the most important coffee‐producing states in Brazil. We observed a significant difference in the incubation period between the populations from Minas Gerais and Espírito Santo when 183 single‐pustule isolates were inoculated onto Catuaí Vermelho IAC 44, a susceptible C. arabica cultivar. Variation in aggressiveness components was observed between and within localities. Isolates with longer incubation periods also tended to have longer latent periods, although there was only a low correlation between these two aggressiveness components (r² = 0.34, P = 2.2 × 10^?16). Low‐sporulating isolates also had significantly longer incubation and latent periods. The H. vastatrix population from Minas Gerais and Espírito Santo is structured by the formation of groups of individuals with differential level of aggressiveness. Our results indicate that the variation in aggressiveness of the Brazilian H. vastatrix population may be associated with the geographic coffee‐producing areas.     

Functional response of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) to Aphis gossypii Glover (Homoptera: Aphididae) in the Laboratory

Stage specific functional response of Harmonia axyridis (Pallas) to varying densities of Aphis gossypii Glover was examined in a simplified cucumber leaf arena under laboratory conditions. All stages of H. axyridis were isolated individually for 24 h with different prey densities at 25 °C and a photoperiod of 16:8 (L:D) h. The number of prey consumed by the predator was checked at 3, 6, 12, and 24 h. All stages of H. axyridis showed a Type II functional response. Based on the random predator equation, estimated attack rates of H. axyridis at 24 h were 0.0037, 0.0442, 0.3590, 0.3228, and 0.1456, and estimated handling times were 4.1001, 2.4575, 0.7500, 0.2132, and 0.1708 h for the first, second, third, and fourth instars, and female adult, respectively.     

Symbiont‐mediated chemical defense in the invasive ladybird Harmonia axyridis

The volatile alkylpyrazines methyl‐ and methoxypyrazines (MPs) present in the reflex bleeds of coccinellid beetles such as the harlequin ladybird beetle Harmonia axyridis are important semiochemicals that function in antipredatory defense behavior. Pyrazines have also been coadapted from a primarily defensive role into pheromones that function in intraspecific communication, attraction, and aggregation behavior. However, the biosynthesis of MPs in ladybird beetles is poorly understood. Here, we tested the hypothesis that MPs could be produced by microbial symbionts in H. axyridis, which generates four different MPs. The evaluation of tissue‐specific MP production showed that MP concentrations were highest in the gut tissue and hemolymph of the beetles rather than the fat body tissue as the presumed site of MP biosynthesis. Furthermore, manipulation of gut microbiota by antibiotic‐containing diets resulted in a lower MP content in adult beetles. The analysis of the bacterial community of the digestive tract revealed the presence of bacteria of the genera Serratia and Lactococcus which are reportedly able to produce MPs. In line with the known diet‐dependent production of MP in H. axyridis, we determined that the presence or relative abundance of some of the potential MP producers (Enterococcus and Staphylococcus) is also diet‐dependent. We hypothesize a potential role of the microbiota in MP production in H. axyridis as a possible example for outsourcing the synthesis of ecologically important semiochemicals to its gut bacteria.     

Nitrogen fertiliser affects the functional response and prey consumption of Harmonia axyridis (Coleoptera: Coccinellidae) feeding on cereal aphids

Predator–prey interactions are influenced by nitrogen availability. Wheat (Triticum aestivum cv. Solstice) plants were provided with four levels of nitrogen and examined the responses of coccinellid predator, Harmonia axyridis to cereal aphids, Rhopalosiphum padi and Sitobion avenae. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. In no‐choice feeding trials, H. axyridis consumed more aphids on low fertilised plants, suggesting a compensatory consumption to overcome reduced biomass (lower aphid size). Total biomass devoured by H. axyridis on all nitrogen fertiliser treatments was not statistically different. Logistic regression analysis of the proportion of prey consumed demonstrated that all developmental stages (larval and adult) of H. axyridis exhibited the type II functional response on all nitrogen fertiliser treatments. The rate of successful search (a′) of third and fourth instars and adults were the same across all fertiliser treatments suggesting that nitrogen fertilisation did not affect a′. Maximum handling time for the first instars of H. axyridis on R. padi (3.81 h^?1) and S. avenae (4.59 h^?1) was on the highest nitrogen treatment while minimum handling time was for the adults of H. axyridis on R. padi (0.20 h^?1) and S. avenae (0.20 h^?1) on the lowest nitrogen treatment. Handling time varied at varying fertiliser treatments within all instars and affected the predator's efficiency. The functional response curve, rate of successful search and handling time provide the information needed to understand the predator–prey interaction between H. axyridis and these cereals aphids. This could lead to the development of a better strategy for the biological control of R. padi and S. avenae at any particular level of nitrogen fertiliser regime in the field crops.     

Intercropping oilseed rape as a potential relay crop for enhancing the biological control of green peach aphids and aphid‐transmitted virus diseases

Tobacco viruses transmitted by green peach aphids, Myzus persicae (Sulzer) (Hemiptera: Aphididae), cause severe disease in flue‐cured tobacco, Nicotiana tabacum L. (Solanaceae), in China and throughout the world. Field experiments were conducted in 2016 and 2017 in Longyan City, Fujian Province, China, to determine whether M. persicae and aphid‐transmitted virus diseases are affected by intercropping of oilseed rape, Brassica napus L. (Brassicaceae), in tobacco fields. The results showed that, compared with those in monocultured fields, the densities of M. persicae and winged aphids in intercropped fields significantly decreased in both 2016 and 2017. In particular, the appearance of winged aphids was delayed by ca. 7 days. Moreover, the densities of Aphidius gifuensis Ashmead (Hymenoptera: Aphidiidae), a parasitoid of the aphid, significantly increased in 2016 and 2017. Accordingly, the incidence rates of aphid‐transmitted virus diseases (those caused by the cucumber mosaic virus, potato virus Y, and tobacco etch virus) significantly decreased in the intercropped fields in 2016 and 2017. Tobacco yields and monetary value significantly increased in 2016 (by 10–25 and 14–29%, respectively) and 2017 (by 17–22 and 22–34%, respectively). Consequently, our results suggest that intercropping oilseed rape in tobacco fields is a good approach to regulating and controlling aphids and tobacco mosaic viruses, for example potyvirus, and this intercropping can help control aphid‐transmitted virus diseases in tobacco.     

Exotic ladybirds for biological control of herbivorous insects – a review

Since the late 19th century, exotic ladybirds (Coleoptera: Coccinellidae) have been used extensively for suppressing herbivorous insects of economic importance. In recent decades, the introduction of non‐native biological control (BC) agents has been greatly limited due to the awareness of the potential non‐target effects of introductions. Nonetheless, recent episodes of biological invasions of economically important pests have raised the need to carefully consider whether the expected benefits of pest control go beyond the possible environmental risks of introduction. To better understand the factors that contributed to successful BC programs, here we review the literature behind classical and augmentative BC using exotic ladybirds. Additionally, by means of case studies, we discuss the BC efficacy of selected exotic species, e.g., Coccinella septempunctata L., Harmonia axyridis (Pallas), and Hippodamia variegata (Goeze), and their position within the communities of predators in the introduced areas of USA, Canada, and Chile. In Europe, much of the research on exotic ladybirds has been conducted on the undesired impact of H. axyridis. Therefore, we summarize the risk assessment data for this species and review the field research investigating the ecological impact on European aphidophagous predators. According to the BIOCAT database of classical BC programs, 212 ladybird species belonging to 68 genera have been released in about 130 years of BC activity, with 14.6% of introductions having resulted in partial, substantial, or complete control of the target pest. However, because post‐release evaluation of establishment and BC success has not always been conducted, this rate could underestimate the successful cases. Among other factors, ladybird establishment and pest suppression mostly depend on (1) intrinsic factors, i.e., high voracity, synchronized predator‐prey life cycle, and high dispersal ability, and (2) extrinsic factors, i.e., adaptability to the new environment and landscape composition. This review contributes to improved understanding of ladybirds as exotic BC agents.     

Feeding behavior of aphids on narrow‐leafed lupin (Lupinus angustifolius) genotypes varying in the content of quinolizidine alkaloids

Since the beginning of breeding narrow‐leafed lupins [Lupinus angustifolius L. (Fabaceae)] with a low alkaloid content, susceptibility to several aphid species has increased. Therefore, the probing and feeding behavior of Aphis fabae Scopoli, Aphis craccivora Koch, Acyrthosiphon pisum (Harris), Myzus persicae (Sulzer), and the well‐adapted Macrosiphum albifrons Essig (all Hemiptera: Aphididae) was studied over 12 h on narrow‐leafed lupin genotypes containing varying amounts and compositions of alkaloids. We used the electrical penetration graph (EPG) technique to obtain information on the influence of alkaloid content and composition on the susceptibility to various aphid species. Results indicated that the total time of probing of A. fabae, A. craccivora, A. pisum, and M. persicae increased with a reduced alkaloid content, whereas the alkaloid content had no influence on M. albifrons. Almost all of the individuals (>93%) conducted sieve element phases on the highly susceptible genotype Bo083521AR (low alkaloid content). A reduced occurrence of phloem phases was observed during the 12‐h recording on the alkaloid‐rich cultivar Azuro, especially for A. pisum (37.5%) and A. fabae (55.0%). Furthermore, aphids feeding on genotypes with low alkaloid content had in most cases significantly longer sieve element phases than when feeding on resistant genotypes (Kalya: low alkaloid content, yet resistant; Azuro: high alkaloid content, resistant), whereas M. albifrons showed the longest phloem phase on the alkaloid‐rich cultivar Azuro. As most significant differences were found in phloem‐related parameters, it is likely that the most important plant factors influencing aphid probing and feeding behavior are localized in the sieve elements. The aphids’ feeding behavior on the cultivar Kalya, with a low alkaloid content but reduced susceptibility, indicates that not only the total alkaloid content influences the feeding behavior but additional plant factors have an impact.