Impact of plant architecture and extraguild prey density on intraguild predation in an agroecosystem

Several models and experimental studies conducted in confined environments have shown that intraguild predation (IGP) can modulate population abundances and structure communities. A number of ecological and abiotic factors determine the nature and frequency of IGP. This study examined the effect of plant architecture and extraguild prey density, and their interactions, on the occurrence of IGP between two species of ladybird, Harmonia axyridis (Pallas) and Propylea quatuordecimpunctata L. (both Coleoptera: Coccinellidae). Theoretical concepts predict that IGP levels would increase with a decrease of both extraguild prey density and plant structural complexity. We conducted a factorial experiment in an open soybean field into which coccinellid larvae were introduced in experimental plots for a period of 5 days. We tested two levels of soybean aphid [Aphis glycines Matsumara (Hemiptera: Aphididae)] density, low and high (100 and 1 000 aphids per plot, respectively), and two levels of plant complexity, low (by removing half of the branches from the soybean plants) and high (by leaving plants intact). We used species‐specific molecular markers to detect the presence of P. quatuordecimpunctata in the digestive tract of H. axyridis. Molecular gut‐content analysis of H. axyridis revealed that rates of IGP were higher (20%) at low aphid density than at high aphid density (<6%). Decreased plant complexity did not impact the frequency of IGP. In accordance with existing literature, this study demonstrates that IGP is amplified at low extraguild prey density. We conclude that considering environmental factors, such as extraguild prey density, is crucial to improve our ability to predict the impact of intraguild predation on community structure and, from an applied perspective, biological control.     

Intraguild predation between the aphidophagous ladybird beetles Harmonia axyridis and Coccinella undecimpunctata (Coleoptera: Coccinellidae): The role of intra and extraguild prey densities

Laboratory cages were used to evaluate the influence of extraguild (EGprey) and intraguild prey (IGprey) densities on the direction, symmetry and magnitude of the intraguild predation (IGP) of the aphidophagous Harmonia axyridis Pallas on Coccinella undecimpunctata L. and vice versa. In order to understand the role of competition between IGprey, the experiments included treatments with one IGpredator, with one or four IGprey and EGprey (Aphis fabae Scopoli) ranging from zero to sufficient aphids to satiate the predators for 12, 24 or 48 h. Increases in EGprey and IGprey densities did not alter the direction, but decreased the magnitude and symmetry of IGP. Predation on one individual of IGprey decreased from more than 80%, in the absence of EGprey, to from 6% to 53%, at higher EGprey densities. Decrease in IGP was less when H. axyridis was the IGpredator. Even at high EGprey densities, eggs and 2nd larval stages of C. undecimpunctata were vulnerable to IGP and the level of predation was 40% and 53%. The presence of more than one IGprey increased the magnitude of IGP mainly at EGprey densities sufficient to satiate the predators for 12 and 24 h, suggesting that competition between the IGpredator and IGprey may be one of the processes promoting IGP. These results and those of other authors suggest that H. axyridis has the potential to be an IGpredator, mainly of the most vulnerable stages of IGprey. Thus, H. axyridis may negatively affect the survival of C. undecimpunctata, when these two species exploit the same resources.     

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.     

Alkaloids provide evidence of intraguild predation on native coccinellids by <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in the field

Over a period of less than 5 years, Belgium was thoroughly invaded by the multicolored Asian coccinellid, Harmonia axyridis. At the same time, a decline of some native coccinellid species was observed in tree habitats. One hypothesis about the cause of this decline was that it might have been due to intraguild predation (IGP) by H. axyridis. In natural conditions, IGP between coccinellids can be tracked by using defensive compounds. Exogenous alkaloids in H. axyridis were therefore examined by gas chromatography-mass spectrometry (GC–MS), using individuals sampled from lime trees that were also occupied by other species of coccinellids. Harmonia axyridis was the dominant species at all life stages, in terms of both numbers of sites occupied and local abundance. The GC–MS analysis of H. axyridis larvae revealed traces of exogenous alkaloids from 19 of the 20 sites and, in nine of those 19 sites, more than 30% of the larvae analyzed contained exogenous alkaloids. Three alkaloids were detected: adaline from Adalia spp., calvine from Calvia spp. and propyleine from Propylea quatuordecimpunctata. Predation by H. axyridis on two different coccinellid species was also detected in the same larva, reinforcing the status of H. axyridis as a top predator. A generalized linear model indicated that IGP frequency was positively influenced by two variables: the abundance of extraguild and intraguild prey; and the interaction between these two variables. Our results therefore support the hypothesis that IGP by H. axyridis on native coccinellids in tree habitats has led to the decline of several of these species.     

Intraguild predation of immature stages of British and Japanese coccinellids by the invasive ladybird Harmonia axyridis

Declines in native aphidophages in North America have been linked to intraguild predation (IGP) by the invasive coccinellid Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). It is feared that many British species will face a similar fate following the recent establishment of H. axyridis in the UK. Meanwhile, H. axyridis exists in apparent ecological equilibrium with other members of its guild in Japan. The impact of H. axyridis on British coccinellids is uncertain but intraguild predatory interactions do occur, particularly amongst immature stages. This study investigates IGP between immature stages of H. axyridis and various British and Japanese coccinellids. The only asymmetric IG predator of H. axyridis at first instar was Anatis ocellata (Linnaeus). Harmonia axyridis engaged in symmetric IGP with Coccinella septempunctata Linnaeus, Calvia quatuordecimguttata (Linnaeus), Harmonia quadripunctata (Pontoppidan) and Eocaria muiri Timberlake, but was the asymmetric IG predator of all other species studied. The level of IGP was high between fourth instar larvae, and frequently biased towards H. axyridis, except in the case of A. ocellata, which again was the only IG predator of H. axyridis. In interactions between fourth instar larvae and pre-pupae, IGP was unidirectional towards H. axyridis for all species except A. ocellata, which acted as both IG predator and IG prey. Pupae were better protected against IGP than pre-pupae but most species were still susceptible to attack by H. axyridis, although IGP was symmetric with A. ocellata, and H. quadripunctata pupae were never attacked. The differences in susceptibility of the various species and developmental stages to IGP by H. axyridis are discussed in relation to physical defence structures. We find no evidence that Japanese species have superior defences to British ones and suggest that behavioural strategies may enable co-existence in the native range. We discuss the relevance of IGP by H. axyridis to the species it is likely to encounter in Britain.     

Suitability of exotic and native lady beetle eggs (Coleoptera: Coccinellidae) for development of lady beetle larvae

Predation upon lady beetle (Coleoptera: Coccinellidae) eggs in the field is most often instances of egg cannibalism by larvae or adults while the majority of the remaining predation events upon coccinellid eggs is done by other species of Coccinellidae. Thus the recent introduction and establishment of Harmonia axyridis in the US could negatively affect native species of Coccinellidae via egg predation. However, little is known regarding the suitability of interspecific coccinellid eggs as a food source for larval development. In this study, it was found that native first or third instar Coleomegilla maculata and Olla v-nigrum larvae were incapable of surviving to the adult stage when provided solely exotic H. axyridis eggs. In stark contrast, H. axyridis larvae survived equally well when cannibalizing eggs or eating eggs of either native species. When C. maculata and O. v-nigrum were grouped as ‘native’ and compared with the exotic H. axyridis, more native eggs were attacked than exotic eggs and a higher percentage of eggs was attacked by H. axyridis larvae. Native and exotic larvae attacked a similar percentage of native eggs but native larvae attacked significantly fewer exotic eggs than did exotic larvae. These data suggest that H. axyridis may prey upon the eggs of these native species, when encountered in the field, compared with the likelihood of the native species preying upon H. axyridis eggs. Therefore, eggs of the native species C. maculata and O. v-nigrum will continue to be subjected to cannibalism and also to possible predation by other native species and the exotic H. axyridis.     

Influence of plant antibiosis through four trophic levels

Summary The effects of an insect herbivore-resistant soybean genotype on relationships between four levels of a trophic system were examined in the laboratory using both greenhouse and field-grown plants. Pre-imaginal development of the predatory pentatomid, Podisus maculiventris (Say), was affected by soybean antibiosis in a manner similar to that of its lepidopteran prey, Pseudoplusia includens (Walker). Pre-imaginal development time was increased, and cumulative weight gain tended to be reduced on both greenhouse and field-grown resistant foliage; although mortality was increased on greenhouse-grown resistant foliage, it appeared unchanged on field-grown foliage. Reproductive capacity of P. maculiventris reared on P. includens larvae that were fed resistant soybean generally was unaffected, although peak progeny production was delayed and extended slightly. Pre-imaginal development and adult emergence of the egg parasitoid Telenomus podisi Ashmead from eggs of P. maculiventris reared on P. includens larvae that were fed resistant soybean were unaffected. However, the overall reproductive capabilities of this parasitoid were reduced. Our results demonstrate that plant antibiosis can influence the biology of organisms over four trophic levels, thus documenting a relationship heretofore undescribed. Determination of the final outcome of the interaction between plant resistance and biological control is difficult. The four trophic level model reported in this study further illustrates the complexity which hinhers a general understanding of these interactions.     

Intraguild predation by Harmonia axyridis (Coleoptera: Coccinellidae) on native insects in Europe: molecular detection from field samples

Intraguild predation (IGP) is a potential mechanism of negative effects on native species populations by invasive non‐native species such as Harmonia axyridis. Molecular techniques (polymerase chain reaction) were used to probe for the presence of various insect (coccinellid, syrphid and chrysopid) prey DNA in the guts of 177 H. axyridis larvae field‐collected in England, France, Germany, Slovakia and the Czech Republic in 2010. Three of the four target prey species were detected in the guts of H. axyridis at the following rates: Adalia decempunctata 9.6%, Adalia bipunctata 2.8% and Episyrphus balteatus 2.8%. IGP on Chrysoperla carnea was not detected. IGP detection of at least one target species was made in England, France, Slovakia and the Czech Republic, but not in Germany. These results strengthen the evidence that H. axyridis is a very generalist predator. Results are discussed in the context of their ecological significance.     

Development and application of molecular gut‐content analysis to detect aphid and coccinellid predation by Harmonia axyridis (Coleoptera: Coccinellidae) in Italy

Despite their positive effect in reducing pest populations, exotic generalist predators sometimes become invasive and contribute to the displacement of indigenous species in the same trophic level. Although laboratory experiments have linked intraguild predation (IGP) to these interactions, field evidence and quantification of IGP are still lacking for most systems. The recent establishment of the exotic Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) in Italy raises concern about the detrimental effect that the ladybird could have on native coccinellids. Here we assessed, under laboratory conditions, the acceptability and suitability of eggs of 2 native ladybirds, Adalia bipunctata L. and Oenopia conglobata (L.), as prey items for H. axyridis larvae. Then we developed primers for molecular gut‐content analysis to detect predation by H. axyridis on the 2 ladybirds and on the aphid Eucallipterus tiliae L. Species‐specific 16S primers were developed for the 3 species and laboratory feeding trials were conducted to quantify the rate of prey DNA breakdown in the gut of H. axyridis. Moreover, to field evaluate primers, H. axyridis 4th instars (n = 132) were systematically collected from linden trees in northern Italy and screened for the presence of prey DNA. Seventy‐three percent and 7% of field collected H. axyridis were positive for aphid and coccinellid DNA, respectively. Predation upon aphid and A. bipunctata was lower than predicted if density dependent consumption was expected, while predation upon O. conglobata was significantly higher. Here, we provided the first evidence of IGP among feral populations of H. axyridis and indigenous ladybird beetles, occurring in Italy.     

Predation by Podisus maculiventris (Hemiptera: Pentatomidae) on Plutella xylostella (Lepidoptera: Plutellidae) larvae parasitized by Cotesia plutellae (Hymenoptera: Braconidae) and its impact on cabbage

Biological control offers potentially effective suppression of the diamondback moth (DBM), Plutella xylostella, a serious pest of Brassica crops. Little is known of whether multiple natural enemies have additive, antagonistic, or synergistic effects on DBM populations. No-choice and choice tests were conducted to assess predation by Podisus maculiventris on DBM larvae parasitized by Cotesia plutellae and unparasitized larvae. In no-choice tests, P. maculiventris preyed on greater numbers of parasitized than unparasitized larvae and greater numbers of young larvae than old larvae. In choice tests with early third instar DBM, there was no difference in predation between parasitized or unparasitized larvae. However, in choice tests with older prey, P. maculiventris preyed on more parasitized than unparasitized larvae. Two field studies were conducted to test if this predator and parasitoid have additive, antagonistic or synergistic effects on DBM populations and plant damage in cabbage (Brassica oleracea var. capitata). In 2002, DBM populations were significantly lower in the presence of C. plutellae but not in the presence of P. maculiventris. There was not a significant interaction between the natural enemies. Plant damage was reduced only with C. plutellae. In 2003, DBM populations were significantly lower in the presence of C. plutellae and P. maculiventris, although the combination of natural enemies did not lead to a non-additive interaction. Plant damage was unaffected by the presence of either natural enemy. Because of its greater predation on parasitized larvae, P. maculiventris could be an intraguild predator of C. plutellae. Yet, their overall combined effect in the field was additive rather than antagonistic.     

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.     

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.     

Density of an intraguild predator mediates feeding group size, intraguild egg predation, and intra- and interspecific competition

Intraguild predation (IGP) is common in most communities, but many aspects of density-dependent interactions of IG predators with IG prey are poorly resolved. Here, we examine how the density of an IG predator can affect feeding group size, IG egg predation, and the growth responses of IG prey. We used laboratory feeding trials and outdoor mesocosm experiments to study interactions between a social intraguild predator (larvae of the wood frog; Rana sylvatica) and its prey (spotted salamander; Ambystoma maculatum). Larvae of R. sylvatica could potentially affect A. maculatum by consuming shared larval food resources or by consuming eggs and hatchlings. However, successful egg predation requires group feeding by schooling tadpoles. We established from five to 1,190 hatchlings of R. sylvatica in mesocosms, then added either 20 A. maculatum hatchlings to study interspecific competition, or a single egg mass to examine IGP. Crowding strongly suppressed the growth of R. sylvatica, and IGP was restricted to the egg stage. In the larval competition experiment, growth of A. maculatum was inversely proportional to R. sylvatica density. In the predation experiment, embryonic mortality of A. maculatum was directly proportional to the initial density of R. sylvatica and the mean number of tadpoles foraging on egg masses. IGP on eggs reduced A. maculatum hatchling density, which accelerated larval growth. Surprisingly, the density of R. sylvatica had no overall effect on A. maculatum growth because release from intraspecific competition via egg predation was balanced by increased interspecific competition. Our results demonstrate that the density of a social IG predator can strongly influence the nature and intensity of interactions with a second guild member by simultaneously altering the intensity of IGP and intra- and interspecific competition.L . A. Burley and A. T. Moyer contributed equally to this work.     

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.     

Intraguild interactions and aphid predators: biological efficiency of Harmonia axyridis and Episyrphus balteatus

Introductions of the harlequin ladybird Harmonia axyridis into Belgium for aphid biological control have been followed by declines in native aphid natural enemies. We first examined, in laboratory and field conditions, the impact of larval mobility of either H. axyridis or Episyrphus balteatus, the most abundant native hoverfly in central Europe, on aphid suppression. The hoverfly larvae consumed more aphids (Megoura viciae) in a short term, whereas the ladybird ones were more efficient in a long term. We second investigated the intraguild interactions between H. axyridis and E. balteatus larvae and adults. In larva pairings between the two species in laboratory microcosms containing aphid (M. viciae) infested broad bean (Vicia faba) plants, H. axyridis had an intraguild predation (IGP) advantage over the hoverfly. When conspecific larvae were paired together on aphid‐infested plants, no cannibalism between them was detected. The presence of either H. axyridis or E. balteatus larvae on aphid‐infested plants negatively influenced the ovipositional behaviour of H. axyridis and E. balteatus females; lower numbers of laid eggs were recorded compared to control treatment. Moreover, eggs laid by E. balteatus females were also dropped as victims of predation mainly by H. axyridis larvae. Our results suggest that while the exotic ladybird was more efficient in aphid biological control, larvae and eggs of the native hoverfly species face increased IGP by H. axyridis, which would contribute, as a consequence, to the decline in E. balteatus population following invasion.     

Direct and indirect effect of cannibalism and intraguild predation in the two sibling Harmonia ladybird beetles

In this study, we focused on the direct (i.e., predation) and indirect (i.e., potential threat from coexisting with a larger individual) effects of cannibalism and intraguild predation (IGP) during larval stages of two sibling ladybird beetles. These effects play an important role in the coexistence of the generalist–common Harmonia axyridis and specialist–rare H. yedoensis (Coleoptera: Coccinellidae). Direct predation effect of cannibalism and IGP was asymmetric in the two sibling ladybird beetles; the fourth instar larvae of H. axyridis were better intraguild predators than cannibals, while the reverse was true in the larvae of H. yedoensis. Neither cannibalism nor IGP significantly affected female body weight in either species. Larval H. axyridis surviving exposure to cannibalism or IGP had a reduced number of ovarioles as adults, whereas adult H. yedoensis ovarioles were not affected. For the indirect effects, longer developmental times in males and females and a lower total number of ovarioles in females were detected in H. axyridis. In H. yedoensis, shorter developmental time of males, lighter adult weight and fewer total ovarioles in females were observed. Olfactometer choice experiments clarified that the fourth instar larvae of H. axyridis avoided the first instar conspecific larvae, while those of H. yedoensis were attracted to the odors from H. axyridis and conspecifics. Thus, H. axyridis has an avoidance mechanism only for cannibalism but not for IGP, whereas H. yedoensis does not have any avoidance mechanism. These different behaviors in the direct and indirect effects of cannibalism and IGP observed in the laboratory may play important roles in the coexistence of generalist–common H. axyridis and specialist–rare H. yedoensis in natural conditions, compensating for the large handicap of H. yedoensis at reproductive interference from H. axyridis.     

Impact of intraguild predation by adult Harmonia axyridis (Coleoptera: Coccinellidae) on Aphis glycines (Hemiptera: Aphididae) biological control in cage studies

The soybean aphid, Aphis glycines Matsumura, has become a principal arthropod pest of soybean in the U.S. since its first detection in 2000. This species threatens soybean production through direct feeding damage and virus transmission. A diverse guild of insect predators feeds on soybean aphid in Michigan including the exotic coccinellid Harmonia axyridis, the native gall midge Aphidoletes aphidimyza and the native lacewing Chrysoperla carnea. In addition to feeding on A. glycines some members of this guild may also engage in intraguild predation. These interactions may produce positive, negative, or neutral impacts on A. glycines biological control. We explored the impact of intraguild predation on soybean aphid population dynamics by comparing aphid populations in microcosms with either A. aphidimyza larvae or C. carnea larvae alone, with both a H. axyridis adult and either A. aphidimyza or C. carnea larvae, and without predators. When H. axyridis was present with larval A. aphidimyza or C. carnea, the lady beetle acted as an intraguild predator. However, intraguild feeding did not result in a release of aphid populations compared with microcosms containing only the intraguild and aphid prey. A similar result was found in field cages. Cages allowing large predators had reduced numbers of A. aphidimyza and C. carnea larvae but also significantly fewer aphids compared with predator exclusion cages. Thus, in both lab and field studies the direct impact of H. axyridis on A. glycines overcame its negative impact as an intraguild predator. Together, these studies indicate that while the exotic H. axyridis does act as an intraguild predator and may contribute to local declines in A. aphidimyza and C. carnea, it is also currently important in overall biological control of A. glycines.     

Evidence for intraguild predation by Podisus maculiventris on a ladybeetle, Coleomegilla maculata: Implications for biological control of Colorado potato beetle, Leptinotarsa decemlineata

The potential for intraguild predation (IGP) between larval and adult life stages of twopolyphagous arthropod predators common in NorthAmerican agroecosystems was studied in thelaboratory. Predators examined were the spinedsoldier bug, Podisus maculiventris Say,and the twelve-spotted ladybeetle, Coleomegilla maculata Lengi. A shared preyitem, eggs of the Colorado potato beetle (CPB),Leptinotarsa decemlineata Say, was alsoprovided to the predators, both to provide analternate food source and to quantify theimpact of IGP on the potential for pestsuppression by these two predators. Experimentswere conducted on single potato leaves inplastic cup arenas, and, subsequently, in cagesenclosing whole potato plants. IGP occurredasymmetrically, with P. maculiventrisadults and nymphs only attacking C. maculata larvae. Even though ladybeetle adultswere generally smaller than soldier bug adults,they were never preyed upon. This appears to bethe first documented case of a coccinellidshowing differential larval and adult immunityto attack by a larger invertebrate predator.The impact of IGP, when it did occur, on CPBegg consumption was equivocal. IGP did notconsistently influence levels of predation onthe eggs. Conversely, even when IGP did notoccur, predation on CPB eggs by both predatorstogether did not increase significantly overlevels inflicted by either predator alone. Theimplications for biological control of CPB bythese predators, which are being considered foraugmentative release in potato and tomato cropsin the United States, are discussed.     

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.     

Intraguild predation among Scolothrips longicornis (Thysanoptera: Thripidae), Neoseiulus californicus and Typhlodromus bagdasarjani (Acari: Phytoseiidae) under laboratory conditions

This study was carried out on the ability of predatory thrips Scolothrips longicornis Priesner to feed on 2 phytoseiid species and vice versa. Also the effect of predation of Neoseiulus californicus (McGregor) on Typhlodromus bagdasarjani Wainstein and Arutunjan and vice versa was evaluated. The larvae, prepupae, and pupae of thrips and the eggs, larvae, and protonymphs of phytoseiids were selected as intraguild prey. The intraguild predation (IGP) among S. longicornis and 2 phytoseiid species was unidirectional and in favor of phytoseiids, i.e., S. longicornis was not able to feed on larval stages of 2 phytoseiids. However, N. californicus and T. bagdasarjani fed on the 1st instar larvae (1.39 and 0.80 per day), 2nd instar larvae (0.87 and 0.55 per day), prepupae (0.51 and 0.48 per day), and pupae of thrips (0.51 and 0.49 per day, respectively). Both phytoseiids fed on eggs, larvae, and protonymphal stages of each other. Females of N. californicus consumed more phytoseiid larvae (2.49 per day) than T. bagdasarjani, which consumed 1.08 N. californicus larvae per day. When Tetranychus urticae was presented as an extraguild prey, intensity of IGP between 2 species of phytoseiids and on larval stages of S. longicornis reduced significantly. Therefore, it is concluded that (i) IGP existed among the 3 examined species and lack of feeding of S. longicornis on 2 phytoseiid species can be justified by its feeding type (monophagy), (ii) N. californicus was much more prone to IGP than was T. bagdasarjani.