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.     

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.     

Chemical protection of Calvia quatuordecimguttata eggs against intraguild predation by the invasive ladybird Harmonia axyridis

Intraspecific and interspecific predation of eggs is a well documented phenomenon amongst aphidophagous coccinellids. The invasive species Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) is known to be a top intraguild predator and reported to attack the eggs of many coccinellid species both in a laboratory setting and in the wild. A previous laboratory study highlighted that while many species’ eggs were highly palatable to H. axyridis, the eggs of Calvia quatuordecimguttata (Linnaeus) (Coleoptera: Coccinellidae) appeared to be extremely well protected from attack. Here we present the results of behavioural experiments testing the hypothesis that substances on the egg surface are responsible for this protection, and report preliminary results of GC-MS analysis of these compounds. When the coatings of C. quatuordecimguttata eggs were removed using hexane, they became significantly more susceptible to predation by neonate H. axyridis larvae. However, their overall palatability was not affected, in that complete consumption was never or rarely observed. This suggests that the surface compounds are a true indicator of unpalatability in this species. The effect of hexane-washing on already palatable conspecific eggs was also analysed but had no significant effect on the susceptibility of eggs to cannibalism. We conclude that the eggs of at least one European species are effectively protected by surface deterrents from intraguild predation by H. axyridis. This effect might be due to both the diversity and abundance of hydrocarbons present within the egg coating, the presence of alkenes and/or the presence of patches of a red substance on the eggs’ surface, which is thought to belong to the acid group. In conjunction with data on the susceptibility of other immature stages of C. quatuordecimguttata, this finding may indicate a decreased risk of the species falling victim to invasive H. axyridis, despite their coincident habitat ranges.     

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.     

Harmonia axyridis: What will stop the invader?

In recent years Harmonia axyridis (Pallas, 1773) (Coleoptera: Coccinellidae) has become a very popular insect among biological control practitioners and scientists, not only for its potential to be an efficient biological control agent but also because it is considered invasive. Individuals of this species were deliberately introduced into several countries for biological control of different arthropods pests. However the predator itself became an invasive species, affecting the dynamics and composition of several guilds through direct or indirect interactions with established species, including intraguild predation. In this paper we discuss the reasons why the species has a high invasiveness and what are the limits to invasion by this species. It is not clear if the invasiveness of the beetle is linked to its biological, ecological and behavioural abilities, or to other factors such as invasibility and interactions between the invaders, the noninvaders, and the habitat, which may in part explain the reasons of its success and help us to answer the question “what will stop the invader?” We also discuss the reason for the absence of the predator in the Azores islands. Despite the intentional introduction of H. axyridis in the Azores and the high number of individuals released, there are no records of this species in the wild, despite recent extensive sampling effort. In this paper we discuss the reasons for the apparent failure or the delay in establishment of the predator. One factor which may hamper the establishment of H. axyridis in some of the Azores islands is the absence of winter environmental conditions, mainly the temperature which is seldom lower than 12°C, essential for the induction of diapause. The lack of success in the establishment could be also related to functional diversity saturation, that is species saturation and competitive exclusion of H. axyridis by other previously established species may be operating.     

The chemical ecology of <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

We review the chemical ecology of the ladybird beetle Harmonia axyridis from the perspective of its invasiveness and the deleterious effects it exerts in the regions it has colonised. We outline the nature and quantification of its chemical defence, and discuss the protection this provides against natural enemies, particularly intraguild predators. We consider the role of infochemicals in location of prey, intraspecific communication and intraguild interactions. We also discuss the role of prey allelochemicals in relation to H. axyridis extreme dietary generalism. Harmonia axyridis poses a number of practical problems for human health and well-being, including “ladybug taint” wine contamination and problems resulting from large aggregations overwintering in buildings. We consider chemical insights into these issues and, in particular, how attractants and repellents might help manage H. axyridis populations through a push–pull strategy. We conclude by discussing future perspectives for research.     

Invasion of the coccinellid community associated with a tropical annual agroecosystem

1. Detailed understanding of the mechanisms enabling or limiting the impacts of invasive generalist predators is needed. Harmonia axyridis is an invasive generalist coccinellid predator that may have destabilised coccinellid communities worldwide and is an excellent candidate for investigating invasion dynamics.
2. We evaluated four hypotheses: (1) Nine years after being detected in Central Brazil, Ha. axyridis has dominated the coccinellid community. This is facilitated by (2) its broader aphid prey range than the other aphidophagous coccinellids, (3) its superiority as an asymmetrical intraguild predator, and (4) the invasibility of the present coccinellid community.
3. We sampled the invertebrate communities associated with six organic vegetable farms in Brazil during 2017 and 2018, conducted a feeding trial to measure prey niche breadth and overlap, and estimated intra- and interspecific interaction strengths corresponding to first-order Lotka-Volterra parameters to evaluate the strength of intraguild predation and create a community matrix.
4. We found (1) Ha. axyridis comprised ≤8.3% of the coccinellid community. (2) Hi. convergens had the broadest prey niche and dominated the prey niche of Ha. axyridis. (3) We could not determine from the population data if Ha. axyridis was the dominant intraguild predator because its density was too low and constant. (4) The community matrix indicated that the present-day coccinellid community was stable.
5. We conclude Ha. axyridis has not become invasive in Central Brazil, in part because it does not have the broadest prey niche breadth and the present coccinellid community is stable to the present perturbation of Ha. 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.     

Harmonia axyridis invasions: Deducing evolutionary causes and consequences

I consider evolutionary approaches to deducing factors that have made the ladybird beetle Harmonia axyridis such a successful invader, and the contribution that studies of this species in its native range can make. Work aiming to demonstrate which (pre)adaptations have made the species so successful often fails to compare these putative characters with those of other ladybirds. This has led to a tendency for “argument by design”‐type claims on characters widely shared by non‐invasive coccinellids. There is good evidence from genetic studies that evolutionary change occurred in invasive populations, contributing to their success. There is some evidence for subsequent evolutionary change after the establishment of invasive H. axyridis, primarily in the native organisms with which the ladybird interacts. I show here that there appears to have been little adaptation in H. axyridis, over about 20 generations, to the alkaloids of one North American native intraguild prey, the ladybird Coleomegilla maculata. Studies of H. axyridis in its native range are important, as they provide a snapshot of the ancestral ladybird, unobscured by subsequent evolutionary change related to its invasiveness. They provide baseline data about phenomena such as interactions with natural enemies and intraguild predation, and they also can provide pointers as to how H. axyridis might further adapt in the regions it has colonized. Harmonia axyridis represents an ideal opportunity for greater international co‐operation between scientists studying this species in its native range in Asia and scientists studying it in Europe, America and Africa, where it is an invasive exotic.     

Harmonia axyridis: an environmental risk assessment for Northwest Europe

In this paper, we summarize the international situation with respect to environmental risk assessment for biological control agents. Next, we apply a recently designed, comprehensive risk evaluation method consisting of a stepwise procedure to evaluate the environmental risks of Harmonia axyridis in Northwest Europe. This resulted in the very clear conclusion that H. axyridis is a potentially risky species for Northwest Europe, because it is able to establish, it has a very wide host range including species from other insect orders and even beyond the class of Insecta, it may feed on plant materials, it can cover large distances (>50 km per year), it does move into non-target areas, it may attack many non-target species including beneficial insects and insects of conservation concern, its activities have resulted in the reduction of populations of native predators in North America, it is known as a nuisance in North America and recently also in Northwest Europe, and it may develop as a pest of fruit in North America. Considering the H. axyridis case, current knowledge would lead to the conclusion that, although the predator is capable to effectively control several pest species, its risks are manifold and it should, thus, not have been released in Northwest Europe. At the time of the first releases in Nortwest Europe in 1995, the available scientific literature made clear that H. axyridis is a large sized polyphagous predator and has a great reproductive capacity in comparison with other ladybird beetles, and that there was a need to study non-target effects because of its polyphagous behaviour. In retrospect, this information should have been sufficient to reject import and release of this species, but it was apparently ignored by those who considered release of this predator in Northwest Europe. The case of Harmonia releases in Northwest Europe underlines that there is an urgent need for harmonized, world-wide regulation of biological control agents, including an information system on risky natural enemy species.     

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.     

Intraguild predation on the parasitoid Aphidius ervi by the generalist predator Harmonia axyridis: the threat and its avoidance

Interactions between natural enemies can be crucial for determining their overall control of pest species, yet the mechanisms that govern such interactions are often poorly understood. The risk of negative effects such as intraguild predation and the possibility of mitigating such risks are important components for ultimately determining the compatibility of biological control agents. We performed a group of experiments to determine whether the coccinellid Harmonia axyridis Pallas (Coleoptera: Coccinellidae) poses an intraguild threat to the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae) and to see whether A. ervi is able to avoid predation by responding to the chemical tracks deposited by H. axyridis. We show that although H. axyridis does not readily consume A. ervi mummies, it preferentially consumes parasitized aphids over unparasitized aphids. We also show that A. ervi can defend against this threat by avoiding oviposition in the presence of H. axyridis chemical tracks. Aphidius ervi parasitized far fewer pea aphids Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae) on plants [Vicia faba L. (Fabaceae)] with H. axyridis chemical tracks in a no‐choice environment at a single‐plant scale. Similarly, when parasitoids could move freely between plants with and without tracks, A. ervi parasitism was higher on plants without tracks. Behavioral observations of A. ervi foraging suggested that this might be because of reduced A. ervi attack rates and patch residence times in the presence of H. axyridis tracks. Despite a risk of intraguild predation by H. axyridis, our study suggests that A. ervi may be able to mitigate this risk by altering its behavior in response to chemical cues.     

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.     

Living with the enemy: parasites and pathogens of the ladybird <Emphasis Type="Italic">Harmonia axyridis</Emphasis>

Harmonia axyridis is an invasive alien predator in many countries across the world. The rapid establishment and spread of this species is of concern because of the threat it poses to biodiversity as a generalist predator. Understanding the mechanisms that contribute to the success of this species as an invader is not only intriguing but also critical to our understanding of the processes governing such invasions. The enemy release hypothesis (ERH) could explain the rapid population growth of many invasive alien species. However, empirical evidence in support of the ERH is lacking. An alternative hypothesis that could explain rapid population growth is evolution of increased competitive ability (EICA). Here we provide an overview of the parasites and pathogens of coccinellids with a particular focus on H. axyridis as a host. We examine the differential susceptibility of host species and highlight the resilience of H. axyridis in comparison to other coccinellids. We recognise the paucity and limitations of available information and suggest that studies, within a life-table framework, comparing life history traits of H. axyridis in both the native and introduced ranges are necessary. We predict that H. axyridis could benefit from both enemy release and EICA within the introduced range but require further empirical evidence.     

Long handling times of Praon unicum by Harmonia axyridis result in protection from predation

The intraguild predator Harmonia axyridis has a longer handling time of Praon unicum mummies in contrast to Aphidius matricariae mummies and the pest aphid Myzus persicae. In addition, the rejection rate of P. unicum is higher as compared to the A. matricariae and M. persicae. Harmonia Axyridis also has a shorter residence times when foraging in P. unicum patches. The longer handling time may provide P. unicum with a refuge from intraguild predation by H. axyridis. Thus, Praon unicum could be a better biocontrol agent than A. matricariae in the presence of intraguild predation, as it will face lower predation rates.     

Do microsporidia function as “biological weapon” for Harmonia axyridis under natural conditions?

Invasive alien species, such as the multicoloured Asian ladybird Harmonia axyridis, are often regarded as major drivers of biodiversity loss. Therefore understanding which characteristics or mechanisms contribute to their invasive success is important. Here the role of symbiotic microsporidia in the hemolymph of H. axyridis was investigated in the context of intraguild predation between wild‐caught H. axyridis and the native ladybird species Coccinella septempunctata. The microsporidia were recently discussed to contribute to the unpalatability of Harmonia for other coccinellids during intraguild predation and to function as “biological weapons”. In the present study, visual detection of microsporidia in hemolymph samples revealed that 73.5 % of H. axyridis were infected. Intraguild predation experiments between larvae of the two species showed a significant competitive advantage for H. axyridis, even against larger larvae of C. septempunctata. Adult C. septempunctata always killed and fed on H. axyridis larvae. However only 11.4 % (4 of 47) of C. septempunctata that fed on infected H. axyridis died within 4 months. In contrast to previous studies this suggests that microsporidia or harmonine, the chemical defense compound of H. axyridis, do not lead to death of C. septempunctata preying on larvae of H. axyridis. Instead our results support the idea that competitive advantage during intraguild predation greatly facilitates the success of H. axyridis and that this may help this highly invasive species to outcompete native species. The impact of microsporidia on Harmonia itself as well as on interspecific interactions require further studies.     

Tawny owl vocal activity is constrained by predation risk

The vocal behaviour of birds may be influenced by many factors, including the risk of being detected by a predator. In Doñana Protected Area, the tawny owl co‐exists alongside its intraguild predator, the eagle owl Bubo bubo. We considered four scenarios to study the vocal behaviour of tawny owls at dusk by analysing: A) the calling rate of all males in 29 sites; B) the calling rate at dusk of males living within the home range of the intraguild predator; C) the calling rate of males living within the home range of the intraguild predator between 60 and 90 min after sunset; and D) the duration of male vocal bouts in visits where eagle owls have called. In scenario A we found that only the number of conspecific males affected the calling rate of tawny owls. In scenario B we observed that the presence of an eagle owl calling constrained the calling rate of the intraguild prey. In scenario C we found that this effect seemed mostly associated to a contemporaneous detection of the intraguild predator’s calls. Finally, in scenario D we found no significant effects on bout duration. These results seem to indicate that tawny owls use their intraguild predator’s calls as a cue to assess predation risk, and then adjust their vocal behaviour in order to minimize predation risk by a predator that may locate its prey by its vocalizations.     

Intraguild responses of aphid predators on apple to the invasion of an exotic species, Harmonia axyridis

The effects of the invasion ofan exotic predator, Harmonia axyridis(Pallas) (Coleoptera: Coccinellidae), wereinvestigated using three experiments on theecology of aphid predators on apple. Oneexperiment, 1992, was collected prior to theH. axyridis invasion, and two others,1996 to 1997 and 1999 to 2000, were collectedafter the invasion. Except for one year, 1999,H. axyridis was the dominant coccinellid,replacing the formerly dominant Coccinellaseptempunctata L. (Coleoptera: Coccinellidae),another exotic species. The dominance of H. axyridis was greater among larvae thanamong adults. There was no apparent effect ofthe H. axyridis invasion on abundance ofthe predator, Aphidoletes aphidimyza(Rondani) (Diptera: Cecidomyiidae) and apossible positive effect on the abundance ofchrysopids (Neuroptera: Chrysopidae). Principal component analysis indicated thatalthough individual species were affected, theoverall effect of H. axyridis invasion onthe predator guild as a whole was negligible. The data indicate that the interaction betweenthe two exotic species, H. axyridis andC. septempunctata, may be allowing nativecoccinellids to become more abundant on applethan when C. septempunctata was thedominant coccinellid.     

Predator biodiversity increases the survivorship of juvenile predators

When predator biodiversity strengthens herbivore suppression, the pattern generally is attributed to interspecific complementarity. However, the relaxation of intraspecific interference within diverse communities has received less attention as an underlying factor, and most experiments to date span much less than one predator generation. Here, working with a community of aphid predators, we compared the survivorship of juvenile predators embedded within diverse versus single-species communities of adult predators. We found that greater predator diversity improved juvenile survivorship for three of four predator taxa (the lady beetles Hippodamia convergens and Coccinella septempunctata, and the bug Nabis alternatus; but not the small bug Geocoris bullatus), whereas survivorship was relatively low when juveniles foraged among only conspecific adults. When aphid densities differed they were lowest for the diverse treatment, and so resource availability could not explain differences in juvenile survivorship. Instead, feeding trials indicated that cannibalism generally posed a greater risk to juveniles than did intraguild predation (with Geocoris again the exception). Our results suggest that the dilution of intraspecific interference may play an important, and perhaps underappreciated, role in shaping predator diversity effects. Furthermore, relatively strong cannibalism but weak intraguild predation has the potential to project diversity effects forward into subsequent generations.     

Ecology of <Emphasis Type="Italic">Harmonia axyridis</Emphasis> in natural habitats within its native range

Originally distributed in northeast Asia, Harmonia axyridis (Coleoptera: Coccinellidae) is now found throughout much of the temperate zone. In its native area, H. axyridis maintains stable populations in heterogeneous and temporary habitats because of its great ability to find prey and reproduce, coupled with density-dependent and self-regulatory population regulation. A negative correlation of H. axyridis on the biodiversity of the aphidophagous community has been observed in its native range. The decrease in biodiversity may be mainly caused by the wider range of habitats available to H. axyridis than to the coexisting species. From a theoretical perspective, density-dependent regulation of H. axyridis populations, e.g., cannibalism, may be more important in maintaining the H. axyridis-dominated system, probably than is intraguild predation. Habitat heterogeneity may also be important to the coexistence of H. axyridis and other predators in both native and invaded areas.