Effect of plant nutrition on aphid size,prey consumption,and life history characteristics of green lacewing

Plant quality can directly and indirectly affect the third trophic level. The predation by all the instars of green lacewing, Chrysoperla carnea （S.） （Neuroptera： Chrysopidae） on the cereal aphids, Rhopalosiphum padi （L.）, and Sitobion avenae （F.） at varying nitrogen fertilizer levels was calculated under laboratory conditions. Wheat plants were grown on four nitrogen fertilizer levels and aphids were fed on these plants and subsequently offered as food to the C. carnea. Aphid densities of 10, 30, and 90 were offered to first, second, and third instar larvae of green lacewing. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. Aphid consumption by green lacewings was reduced with the increase in nitrogen content in the host plants of aphids. Predation of both aphid species by first, second, and third instars larvae of C. carnea was highest on aphids reared on plants with the lowest rate of fertilization, suggesting a compensatory consumption to overcome reduced biomass （lower aphid size）. Total biomass devoured by C. carnea on all nitrogen fertilizer treatments was not statistically different. Additionally, the heavier host prey influenced by the plant nutrition had an effect on the life history characteristics of green lacewings. The larval duration, pupal weight, pupal duration, fecundity, and male and female longevity were significantly affected by the level of nitrogen fertilization to the aphid＇s host plants, except for pupal duration when fed on S. avenae. This study showed that quantity of prey supplied to the larvae affects the prey consumption and thereafter the life history characteristics of green lacewings.     

The influence of nitrogen fertiliser on the population development of the cereal aphids Sitobion avenae (F.) and Metopolophium dirhodum (Wlk.) on field grown winter wheat

The effect of nitrogen fertiliser on populations of the cereal aphids Sitobion avenae and Metopolophium dirhodum on winter wheat was investigated in a three year field experiment. Naturally occurring aphid populations were monitored on three nitrogen treatments; none, nitrogen application using Canopy Management guidelines (130–210 kg ha^-1) and conventional practice (190 kg ha^-1). Inoculations of laboratory reared S. avenae were used to enhance field populations on half the plots. Natural populations of M. dirhodum remained below the current UK spray threshold level of two-thirds of shoots infested at the start of flowering, or five aphids per shoot in all years, whilst populations of S. avenae exceeded the threshold in all years. The response of the two species to nitrogen differed. Significantly higher populations of M. dirhodum were recorded in both treatments which received nitrogen in all years, whilst the response of S. avenae varied between years. In 1994 and 1995 when environmental conditions favoured aphid development, higher populations were recorded in the two treatments which received nitrogen. In 1993 when high rainfall created unfavourable conditions, higher populations were recorded in the plots receiving no nitrogen. Differences in peak density and cumulative aphid index of S. avenae resulted from differences in the rate of population increase between ear emergence and peak density on the different treatments. Populations prior to ear emergence were higher in the plots which received nitrogen but the differences were not statistically significant. There was no evidence of a difference in the timing of population decline in the different treatments. In 1993 higher levels of infection by entomopathogenic fungi were observed in all treatments. Significantly higher levels of infection were recorded in the treatments receiving nitrogen, which may have accounted for the lower S. avenae populations recorded. It is possible that the larger canopies recorded in these treatments produced conditions which favoured infection by fungi, thereby limiting aphid population growth. The results indicate that application of nitrogen increases natural populations of M. dirhodum, and under favourable conditions, populations of S. avenae. However, in suboptimal climatic conditions, the application of nitrogen fertiliser can lead to lower populations of 5. avenae. The data also suggest that there is no consistent difference between a conventional and Canopy Managed approach to nitrogen fertiliser use in terms of the risk of infestation by cereal aphids.     

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.     

The effect of past natural enemy activity on host‐plant preference of two aphid species

The selection of a host of high nutritional quality is of great importance to the development of offspring of larvipositing aphids, as is the avoidance of natural enemies. Little is known, however, about their ability to select host plants based on these factors. This article tests the preference of aphids Sitobion avenae (Fabricius) and Rhopalosiphum padi (L.) (both Hemiptera: Aphididae) for different winter wheat cultivars, Triticum aestivum (L.) (Poaceae), and their ability to detect and avoid predators in sacrifice of their most preferred host. In both species a preference was observed for nutritionally superior hosts. The preference of both species then exhibited a change towards a nutritionally inferior host after infestations of the harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), that had been consuming conspecific aphids. This investigation opens the door to the interesting prospect of the ability of aphids to make complex decisions regarding a compromise between high‐quality nutrition and avoidance of predation.     

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.     

Lethal and behavioral sublethal side effects of thiamethoxam on the predator Harmonia axyridis

Thiamethoxam (TMX) is one of the most effective neonicotinoid insecticides for the control of green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), and various side effects can be expected in its natural enemies. The multicolored Asian lady beetle or harlequin ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), is an important predator against M. persicae in greenhouses and fields. In this study, we evaluated the toxicity of TMX to H. axyridis and the effect on the functional response of this predator to M. persicae through three routes of exposure. Acute toxicity bioassays indicated that LC₅₀ values of TMX on H. axyridis through direct residual contact (‘contact’), systemic application (‘systemic’), and leaf‐dip treatment (‘leaf‐dip’) were 18.99, 21.26, and 15.39 mg a.i. l^?1, respectively. The hazard quotient indicated a potential hazard of this agrochemical regardless of the exposure routes. The mortality caused by the lowest rate, 2 mg a.i. l^?1, was not significantly different compared with the control group. For the three routes of sublethal TMX exposure, the type‐II functional response was a good fit to the prey consumption of H. axyridis. Predation was most affected by leaf‐dip treatment, followed by contact and systemic treatments, which had similar effect. For all exposure routes, the predation capacity of the predator recovered quickly after transfer to untreated cabbage leaves. Thiamethoxam applied systemically was the least toxic to H. axyridis and did not affect the functional response of the predator. However, the sublethal effects of TMX through both contact and leaf‐dip application may reduce the population growth of H. axyridis and consequently impair the biological control of M. persicae by this predator. These results illustrate that the assessment of potential effects of TMX on H. axyridis is crucial to develop effective integrated pest management programs for M. persicae in China.     

Analysis of the aggregation behavior in the larvae ofHarmonia axyridis Pallas (Coleoptera: Coccinellidae) to prey colony

Summary The searching and handling behaviors ofHarmonia axyridis larvae to the colony ofRhopalosiphum padi were experimentally examined and the processes of their aggregation to the prey colony was analyzed. All the instar larvae searched for the prey at random and they have no preference to the prey colony, but except the 1st instar they tend to aggregate to the plants with prey colonies. The 1st instar larvae tend to stay on the plants they once located. The 2nd to 4th instar larvae often emigrate from the plants without prey colony but seldom emigrate from the plants with prey colonies, and consequently, they aggregate to the plants with prey colonies. The expense of time to eat prey (in the 2nd and 3rd instars) and the change of searching behavior for the prey after feeding (in the 3rd and 4th instars) are responsible for the larval concentration to prey colony as a trapping effect for predators to prey colony.     

Interaction between Podisus maculiventris and Harmonia axyridis two predators used in augmentative biological control in greenhouse crops

Intraguild predation (IGP) betweenthe pentatomid Podisus maculiventris(Say) and the coccinellid Harmoniaaxyridis (Pallas) in the absence or presenceof the extraguild prey Spodopteralittoralis (Boisduval) and Myzuspersicae (Sulzer) was studied in thelaboratory. Interactions were asymmetric infavor of the pentatomid. Podisusmaculiventris readily fed upon eggs and larvaeof H. axyridis, but adult beetles wererarely attacked. Success of attacks by P.maculiventris was stage dependent, fourthinstars and adults being more successful inkilling ladybeetle larvae than second instars.Attacks by H. axyridis on the pentatomidwere rare and none of them were successful. Theeffect of introducing extraguild prey on thelevel of IGP was tested both in petri dishesand on caged sweet pepper plants. Whensufficient numbers of S. littoralislarvae were present to satiate the pentatomid,predation on H. axyridis larvae decreasedsignificantly, indicating that the coccinellidis a less preferred or less vulnerable prey.When the aphid M. persicae was presentedas extraguild prey, levels of IGP were notaltered. Nymphs of P. maculiventrissuccessfully completed development whenexclusively fed on larvae of H. axyridis,but developmental time was longer than onlepidopteran prey. No pentatomid nymphs reachedadulthood on aphids alone. IGP by P.maculiventris on H. axyridis may be ofsome importance in greenhouse crops, where bothpredators are being used increasingly inaugmentative biological control programs.Nonetheless, it is expected that in practicelarger larvae and adults of H. axyridiswill escape most attacks by the pentatomid.     

Prey capture performance in hatchlings of two sibling Harmonia ladybird species in relation to maternal investment through sibling cannibalism

1. To elucidate the factors responsible for the evolution of host specialisation, prey capture performance in hatchlings of two sibling ladybird species, Harmonia yedoensis and H. axyridis, feeding on four aphid species was examined. Harmonia yedoensis is a specialist predator that preys only on pine aphids in the field, whereas H. axyridis is a generalist predator with a broad prey range. 2. In H. yedoensis, sibling cannibalism in each clutch was intense and predation against pine aphid as well as other aphid species was moderately successful. In contrast, the predation success rate of H. axyridis against pine aphid was quite low. 3. Moreover, it was experimentally shown that increased maternal investment enhanced prey capture performance against pine aphid in H. yedoensis but not in H. axyridis hatchlings, despite their increased body size due to maternal investment. 4. In addition, morphological and behavioural analysis showed that hatchlings of H. yedoensis had longer legs and a larger head capsule size and could walk faster than H. axyridis. 5. In summary, the interactive effects between a large amount of maternal investment and morphological specialisation of the first instars may enable H. yedoensis to capture the pine aphid efficiently, a highly elusive prey for ladybird hatchlings. The ability of H. yedoensis to utilise the three other aphid species in addition to the pine aphid suggests that a trade‐off in prey capture performance is not the main factor in the host specialisation of H. yedoensis.     

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.     

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.     

Performance of clones and morphs of two cereal aphids on wheat plants with high and low nitrogen content

The great variability of the aphid life cycle and their tendency for host alternation gives rise to aphid clones and morphs. Inter‐ and intraclonal variability may be observed in the responses of aphids to various environmental factors. In this study we aimed to evaluate the influence of intrinsic factors (clone and morph type) on the intrinsic rate of increase (r_m) of the English grain aphid, Sitobion avenae (Fabricius), and the bird cherry‐oat aphid, Rhopalosiphum padi (Linnaeus). For each species four apterous clones were collected from established laboratory colonies and compared to assess their relative fitness on high‐ and low‐nitrogen wheat plants under laboratory conditions. The clones had significantly different intrinsic rates of increase on high‐ and low‐nitrogen plants. All R. padi clones had a higher intrinsic rate of increase and mean relative growth rate than S. avenae. Experiments were also conducted to compare the mean fecundity of apterous and alate morphs of S. avenae and R. padi clones on high‐ and low‐nitrogen wheat plants. On high‐nitrogen plants the apterous morphs of S. avenae clones had significantly higher mean fecundity than alate morphs. There were no significant differences between the mean fecundity of alate morphs of the same species on high‐ and low‐nitrogen plants. The results support the idea of better fitness of specific clones/morphs on certain host plants due to higher and lower intrinsic rates of increase.     

Behavioral Response to Larval Tracks and the Influence of Tracks on Intraguild Scavenging by Coccinellid Larvae

A paired design was used to determine that Harmonia axyridis 4th instars were not influenced by the presence of conspecific larval tracks, but well-fed H. axyridis 4th instars spent less time on plants that contained tracks left by Coleomegilla maculata 4th instars. To determine if the presence of larval tracks influences intraguild scavenging by H. axyridis 4th instars, dead 4th instars were placed in Petri dishes that contained or did not contain larval tracks. The presence of larval tracks did not influence the feeding frequency or the amount of time before feeding. However, larvae dragged their pygopod on dish surfaces more frequently if the dish contained larval tracks. In addition, starved H. axyridis larvae were more likely to feed on the prey and dragged their pygopod less frequently than well-fed larvae.     

Intraguild predation on the aphid parasitoid <Emphasis Type="Italic">Aphelinus asychis</Emphasis> by the ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

We investigated intraguild predation (IGP) on an aphid parasitoid, Aphelinus asychis Walker (Hymenoptera: Aphelinidae), by the multicolored Asian ladybird, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), and used the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae) as the prey/host in the laboratory. The ladybirds reared on artificial diet and on aphids consumed more aphids than mummies, while those reared on parasitized aphids consumed similar numbers of aphids and mummies. The ladybirds chose more mummies in treatments when mummies were more abundant, and more aphids when numbers of aphids and mummies were equal, or when aphids were more abundant. However, at all density treatments, rejection rates of mummies (36%) were much greater than of aphids (2%). H. axyridis prey on more aphids than A. asychis mummies, which enhances biological control by the two species. However, prior feeding experience affected subsequent choice, increasing the competition between natural enemies which would reduce their combined effectiveness for biological control.     

The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Possible coexistence of Harmonia axyridis Pallas (Coleoptera: Coccinellidae) and Diaeretiella rapae M'Intosh (Hymenoptera: Braconidae) in the biological control of Lipaphis erysimi (Homoptera: Aphididae)

To study the interactions between the aphidophagous predator Harmonia axyridis Pallas (Coleoptera: Coccinellidae) and the specialist aphid parasitoid Diaeretiella rapae M'Intosh (Hymenoptera: Braconidae) in the biological control of mustard aphid, Lipaphis erysimi (Homoptera: Aphididae), the prey discrimination by H. axyridis among unparasitized, non-mummified parasitized and mummified aphids was examined under laboratory conditions. Prey/host selections were also tested by offering L. erysimi at various developmental stages to assess the possibility of coexistence between the two species, so the prey preference of H. axyridis when D. rapae parasitize aphids, and the host preference of D. rapae when H. axyridis interfered with the parasitization were detected. We found that H. axyridis could discriminate against mummies rather than non-mummified parasitized aphids. The ladybug showed a significantly positive preference for adult prey when D. rapae turned aphids into mummies, while D. rapae tended to parasitize younger nymphal aphids when H. axyridis was introduced. The present study suggests the prey discrimination against mummies by H. axyridis, and indicates that H. axyridis and D. rapae can avoid resource competition by attacking different and non-overlapping developmental stages of aphid. Thus, H. axyridis and D. rapae can potentially coexist and establish a stable ecosystem in the biological control of L. erysimi.     

Effect of resource gradient on age and size at maturity and their influence on early‐life fecundity in the predatory Asian lady beetle,Harmonia axyridis

Variation in food availability impacts the performance of insects in terms of their size and age to maturity and fecundity. Age at maturity determines how quickly individuals in a population can start to reproduce and how much they can reproduce. Results from studies on various insect species show that food availability influences the size and fecundity of adult females. It is predicted that under poor growth conditions, variation in size is low, but variation in age at maturity is considerable. This prediction was examined in a widely distributed lady beetle species, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), a predator of aphids and coccids. Using a food gradient from low to high aphid prey density, performance of females that were reared on excess food was recorded for pre‐reproductive duration, size at reproductive maturity, number of aphids consumed, and fecundity in the first 10 days of their reproductive period. Results suggested that female H. axyridis that were reared on surplus food when kept at low prey density (poor growth condition) took, on average, three times longer to attain maturity and produced, on average, 14 times fewer eggs than females that were also reared on surplus food, but kept at high prey density (good growth condition). Females performed best at a prey density of 30 aphids per female per 150 cm². Results suggested that the current food availability significantly influenced the age and size of females at maturity and their fecundity. Age and size at maturity of female lady beetles showed non‐linear responses to prey density as well as the occurrence of a minimum size of females, below which H. axyridis females fail to mature. The steep slope recorded at lower prey densities suggests relatively high variation in age at maturity but low variation in size.     

Effects of mummy consumption on fitness and oviposition site selection on Harmonia axyridis

Intraguild predation (IGP) has been commonly reported between predators and parasitoids used as biological control agents as predators consuming parasitoids within their hosts. However, the effect of parasitoid–mummy consumption on the fitness of the predator and subsequent oviposition site selection have not been well studied. In our study, we conducted two laboratory experiments to examine the influence of Aphidius gifuensis Ashmead (Hymenoptera: Braconidae) mummies as prey on fitness and subsequently oviposition site selection of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Results indicate that when H. axyridis was reared on A. gifuensis mummies only, its larval development was prolonged, and body weight of the 4th instar larvae and newly emerged adults, and fecundity decreased. Moreover, H. axyridis did not exhibit oviposition preference on plants infested with unparasitized aphids or aphids parasitized for shorter than 9 days. However, compared with plants with mummies (parasitized ≥9 days), H. axyridis laid more eggs on plants with unparasitized aphids. In contrast, H. axyridis previously fed with A. gifuensis mummies did not show a significant oviposition preference between plants with unparasitized aphids and those with mummies (parasitized ≥9 days). Overall, our results suggest that mummy consumption reduced the fitness of H. axyridis. Although H. axyridis avoided laying eggs on plants with A. gifuensis mummies, prior feeding experience on A. gifuensis mummies could alter the oviposition site preference. Thus, in biological control practice, prior feeding experience of H. axyridis should be carefully considered for reduction of IGP and increase of fitness of H. axyridis on A. gifuensis.     

A sublethal effect on native Anthocoris nemoralis through competitive interactions with invasive Harmonia axyridis

1. The invasive ladybird Harmonia axyridis Pallas (Coleoptera: Coccinellidae) is implicated in declines of native ladybirds with intraguild predation identified as an underlying mechanism, however, less studied are the effects on non‐coccinellid predators. Intraguild predation between Anthocoris nemoralis Fabricius (Heteroptera: Anthocoridae) and H. axyridis and the relative effects of inter‐ and intraspecific competition on predators on potted Tilia cordata Mill. was investigated, at two aphid densities. 2. Intraguild predation was unidirectional in favour of H. axyridis in Petri dishes although a low level of ladybird egg predation by A. nemoralis occurred. However, on trees intraguild predation was rare. 3. A positive effect of aphid density on H. axyridis weight gain, and increased survival when reared with A. nemoralis on potted trees indicated stronger effects of intraspecific competition on H. axyridis. Consistent suppression of aphids by ladybirds revealed their superior competition for aphids. 4. Anthocoris nemoralis weight gain was reduced when reared with H. axyridis on potted trees, indicating a potential for a sublethal effect through interspecific competition. Survivorship of A. nemoralis did not differ between con‐ and heterospecific treatments, but high aphid density enhanced A. nemoralis survival. Overall, high mortality among nymphs across treatments suggested effects related to both inter‐ and intraspecific competition for resources. 5. Results provide evidence of a sublethal effect by invasive H. axyridis through resource competition, suggesting that where these predators co‐occur, competitive interactions for shared prey are more likely to affect A. nemoralis populations than intraguild predation by H. axyridis.     

Effect of size,extraguild prey and habitat complexity on intraguild interactions: a case study with the invasive ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis> and the hoverfly <Emphasis Type="Italic">Episyrphus balteatus</Emphasis>

Intraguild predation (IGP) is a widespread interaction between predatory arthropods, and is influenced by several factors. The harlequin ladybird, Harmonia axyridis (Coleoptera: Coccinellidae), has frequently been reported as an intraguild predator of other Coccinellidae, but little is known about its interactions with other aphidophagous predators, including syrphids. This study investigated the incidence of IGP between H. axyridis and Episyrphus balteatus (Diptera: Syrphidae), the most abundant hoverfly species in Europe and a commercially available aphid biocontrol agent. The influence of size, presence of extraguild prey and habitat complexity were investigated through laboratory experiments in Petri dishes and on potted broad bean plants. In both types of arenas, IGP between H. axyridis and E. balteatus was found to be asymmetric, with the coccinellid in the majority of cases being the intraguild predator. There was a significant effect of size on the frequency of IGP. The efficiency of H. axyridis as an intraguild predator increased with the developmental stage. Early instars of E. balteatus were the most vulnerable to IGP. Pupae of either species were not attacked. In the presence of extraguild prey, the frequency of IGP was substantially reduced. However, IGP still occurred, mainly in combinations of older larvae of H. axyridis with first or second instars of E. balteatus. The size of the arena affected the incidence of IGP in combinations with second instars of E. balteatus, but not in combinations with third instars. Field research is needed to elucidate the ecological relevance of IGP among these predators.