Der Asiatische Marienkäfer als Modell

The harlequin ladybird as a model in invasion biology The harlequin ladybird Harmonia axyridis has been established as a promined model in invasion biology because it particularly suited to test hypotheses explaining the successful spread of some introduced species. Recent research elucidated that H. axyridis carries parasites which can infect native ladybird species if the feed on its eggs or larvae within intraguild predation. Besides these “bioweapons” H. axyridis accumulates small molecules in its hemolymph which provide as chemical weapons protection against pathogens and predators. The application of state of the art genetic tools will help to elucidate adaptations mediating invasive success at the molecular level.     

异色瓢虫对与本土蚜虫共存的外来扶桑绵粉蚧的搜寻和捕食行为

[目的]外来入侵物种作为猎物可能影响本土广谱捕食者的搜寻和捕食行为.本研究旨在揭示异色瓢虫Harmonia axyridis对与本土豌豆修尾蚜Megoura japonica混合发生的外来入侵扶桑绵粉蚧Phenacoccus solenopsis的搜寻和捕食行为.[方法]试验前对异色瓢虫设两个饲喂猎物种类处理:用扶桑绵...     

Do anuran larvae respond behaviourally to chemical cues from an invasive crayfish predator? A community-wide study

Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators’ diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.     

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

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

     

Predator–prey naïveté, antipredator behavior,and the ecology of predator invasions

We present a framework for explaining variation in predator invasion success and predator impacts on native prey that integrates information about predator–prey naïveté, predator and prey behavioral responses to each other, consumptive and non‐consumptive effects of predators on prey, and interacting effects of multiple species interactions. We begin with the ‘naïve prey’ hypothesis that posits that naïve, native prey that lack evolutionary history with non‐native predators suffer heavy predation because they exhibit ineffective antipredator responses to novel predators. Not all naïve prey, however, show ineffective antipredator responses to novel predators. To explain variation in prey response to novel predators, we focus on the interaction between prey use of general versus specific cues and responses, and the functional similarity of non‐native and native predators. Effective antipredator responses reduce predation rates (reduce consumptive effects of predators, CEs), but often also carry costs that result in non‐consumptive effects (NCEs) of predators. We contrast expected CEs versus NCEs for non‐native versus native predators, and discuss how differences in the relative magnitudes of CEs and NCEs might influence invasion dynamics. Going beyond the effects of naïve prey, we discuss how the ‘naïve prey’, ‘enemy release’ and ‘evolution of increased competitive ability’ (EICA) hypotheses are inter‐related, and how the importance of all three might be mediated by prey and predator naïveté. These ideas hinge on the notion that non‐native predators enjoy a ‘novelty advantage’ associated with the naïveté of native prey and top predators. However, non‐native predators could instead suffer from a novelty disadvantage because they are also naïve to their new prey and potential predators. We hypothesize that patterns of community similarity and evolution might explain the variation in novelty advantage that can underlie variation in invasion outcomes. Finally, we discuss management implications of our framework, including suggestions for managing invasive predators, predator reintroductions and biological control.     

Native predator eats invasive toxic prey: evidence for increased incidence of consumption rather than aversion-learning

Contemporary adaptation of native prey species to invasive predators has been relatively well documented, but that of native predators to invasive prey has received less attention. Because the level of impact an invasive species will have on its predators versus its prey will determine changes in community trophic structure, it is important to understand how native predators respond to novel prey. Here we examine the response of native fence lizards to the invasion of red imported fire ants, a novel toxic prey. Examining invaded and uninvaded lizard populations, we tested whether or not aversion-learning occurs in juvenile fence lizards over successive feedings (within lifetime), how previous fire ant exposure may affect avoidance behavior (over generations), and whether population differences are consistent when prey choice exists. We also examine rates of phenotypic divergence in traits associated with the native species as both predator and prey. Aversion-learning did not occur in either population. Instead, the incidence of fire ant consumption increased over both successive feedings and generations. Lizards from the fire ant invaded population had a higher propensity to eat fire ants than fire ant-naïve lizards, even when given a choice between prey items. We found greater phenotypic divergence in traits associated with the native species as predator on, versus as prey to, fire ants. Although the strategy of eating these novel toxic prey can impose survival costs in the short term, over the longer-term, eating fire ants may cost little or even benefit survivors.     

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.     

Impact of oilseed rape expressing the insecticidal cysteine protease inhibitor oryzacystatin on the beneficial predator Harmonia axyridis (multicoloured Asian ladybeetle)

Insect-resistant transgenic plants have been suggested to have deleterious effects on beneficial predators through transmission of the transgene product by the pest to the predator. To test this hypothesis, effects of oilseed rape expressing the cysteine protease inhibitor oryzacystatin-1 (OC-1) on the predatory ladybird Harmonia axyridis were investigated using diamondback moth Plutella xylostella as the pest species. As expected, oilseed rape expressing OC-1 had no effects on either development or survival of the pest, which utilizes serine digestive proteases. Immunoassays confirmed accumulation of the transgene product in pest larval tissues at levels of up to 3 ng per gut. Characterization of proteolytic digestive enzymes of H. axyridis demonstrated that larvae and adults utilize cysteine and aspartic proteases; the former activity was completely inhibited by oryzacystatin in vitro. However, when H. axyridis larvae consumed prey reared on OC-1 expressing plants over their entire life cycle, no significant effects upon survival or overall development were observed. The inhibitor initially stimulated development, with a shortening of the developmental period of the second instar by 27% (P < 0.0001) accompanied by a 36% increase in weight of second instar larvae (P = 0.007). OC-1 had no detrimental effects on reproductive fitness of adult H. axyridis. Interestingly there was a significant increase in consumption of OC-1 dosed prey. The results show that prey reared on transgenic plants expressing a protein which inhibited ladybird digestive enzymes in vitro had no effects in vivo; the ladybird was able to up-regulate digestive proteases in response to the inhibitor.     

Costs and benefits of defences induced by predators differing in dangerousness

While theoretical studies predict that inducible defences should be fine-tuned according to the qualities of the predator, very few studies have investigated how dangerousness of predators, i.e. the rate at which predators kill prey individuals, affects the strength of phenotypic responses and resulting benefits and costs of induced defences. We performed a comprehensive study on fitness consequences of predator-induced responses by involving four predators (leech, water scorpion, dragonfly larva and newt), evaluating costs and benefits of responses, testing differences in dangerousness between predators and measuring responses in several life history traits of prey. We raised Rana dalmatina tadpoles in the presence of free-ranging predators, in the presence of caged predators, and exposed naive and experienced tadpoles to free-ranging predators. Tadpoles adjusted the intensities of their behavioural and morphological defences to predator dangerousness. Survival was lower in the nonlethal presence of the most dangerous predator, while we could not detect costs of induced defences at or after metamorphosis. When exposed to free-ranging predators, small, but not large, tadpoles benefited from exhibiting an induced phenotype in terms of elevated survival when compared to naive tadpoles, but we did not observe higher survival either in tadpoles exhibiting more extreme phenotypes or in tadpoles exposed to the type of predator they were raised with. These results indicate that while predator-induced defences can mirror dangerousness of predators, costs and benefits do not necessarily scale to the magnitude of plastic responses.     

Behavioral plasticity in an invaded system: non-native whelks recognize risk from native crabs

Inducible defenses have the potential to affect both invasion success and the structure of invaded communities. However, little is known about the cues used for risk-recognition that influence the expression of inducible defenses in invasive prey, because they involve a novel threat. In laboratory experiments, we investigated behavioral defenses induced by a native crab on two invasive oyster drills (marine whelks Urosalpinx cinerea and Ocinebrina inornata). Both drills hid more often and reduced their feeding rates when they detected predators consuming conspecific prey. Examination of the responses of U. cinerea to specific cue sources (predator kairomones, conspecific alarm cues) indicated that this species had the strongest responses to cues from injured conspecifics, but that it did recognize the novel crab predator. Our observation of native predator (per se) recognition by an invasive marine prey is novel. In addition, we observed that neither species of drill reduced their defensive behavior to reflect predation risk shared by a group of prey. The lack of density dependence in risk-assessment could cause populations of invasive prey to transmit both quantitatively and qualitatively different community effects over the course of an invasion as abundance changes. Together, these findings demonstrate several ways that the risk-assessment strategies could be important in establishment and post-establishment dynamics of invasive prey.     

A field demonstration of the costs and benefits of group living to edible and defended prey

Both theoretical and laboratory research suggests that many prey animals should live in a solitary, dispersed distribution unless they lack repellent defences such as toxins, venoms and stings. Chemically defended prey may, by contrast, benefit substantially from aggregation because spatial localization may cause rapid predator satiation on prey toxins, protecting many individuals from attack. If repellent defences promote aggregation of prey, they also provide opportunities for new social interactions; hence the consequences of defence may be far reaching for the behavioural biology of the animal species. There is an absence of field data to support predictions about the relative costs and benefits of aggregation. We show here for the first time using wild predators that edible, undefended artificial prey do indeed suffer heightened death rates if they are aggregated; whereas chemically defended prey may benefit substantially by grouping. We argue that since many chemical defences are costly to prey, aggregation may be favoured because it makes expensive defences much more effective, and perhaps allows grouped individuals to invest less in chemical defences.     

Why aren't warning signals everywhere? On the prevalence of aposematism and mimicry in communities

Warning signals are a striking example of natural selection present in almost every ecological community – from Nordic meadows to tropical rainforests, defended prey species and their mimics ward off potential predators before they attack. Yet despite the wide distribution of warning signals, they are relatively scarce as a proportion of the total prey available, and more so in some biomes than others. Classically, warning signals are thought to be governed by positive density-dependent selection, i.e. they succeed better when they are more common. Therefore, after surmounting this initial barrier to their evolution, it is puzzling that they remain uncommon on the scale of the community. Here, we explore factors likely to determine the prevalence of warning signals in prey assemblages. These factors include the nature of prey defences and any constraints upon them, the behavioural interactions of predators with different prey defences, the numerical responses of predators governed by movement and reproduction, the diversity and abundance of undefended alternative prey and Batesian mimics in the community, and variability in other ecological circumstances. We also discuss the macroevolution of warning signals. Our review finds that we have a basic understanding of how many species in some taxonomic groups have warning signals, but very little information on the interrelationships among population abundances across prey communities, the diversity of signal phenotypes, and prey defences. We also have detailed knowledge of how a few generalist predator species forage in artificial laboratory environments, but we know much less about how predators forage in complex natural communities with variable prey defences. We describe how empirical work to address each of these knowledge gaps can test specific hypotheses for why warning signals exhibit their particular patterns of distribution. This will help us to understand how behavioural interactions shape ecological communities.     

Do exotic generalist predators alter host plant preference of a native willow beetle?

1 Selection can favour herbivores that choose host plants benefitting their offspring either by enhancing growth rates or by increasing larval defences against native predators. For exotic predator species that feed on herbivores, their success with invading new habitats may depend upon overcoming defences used by native prey. Whether exotic predators can alter herbivore host choice has remained unexamined. Therefore, we compared the efficacy of larval defence by Chrysomela knabi (a native beetle species) that had fed on two native willow hosts: Salix sericea (a phenolic glycoside (PG)-rich species) and Salix eriocephala (a PG-poor species), when attacked by exotic generalist predators. In addition, the preference and performance of C. knabi on S. sericea and S. eriocephala was examined.
2  Chrysomela knabi preferred and performed better on S. sericea. In a common garden, adult C. knabi were nine-fold more common and oviposited five-fold more frequently on S. sericea than on S. eriocephala . In the laboratory, adult feeding preference on leaf discs and survival rates of larvae were both greater on S. sericea , and time to pupation was shorter.
3  Chrysomela knabi larvae produced significantly more salicylaldehyde when fed S. sericea leaves than when fed S. eriocephala leaves. Additionally, those larvae with greater salicylaldehyde had reduced predation by two exotic generalist predators, Harmonia axyridis larvae and juvenile Tenodera aridifolia sinensis .
4 The results obtained in the present study suggest that selection favoured the preference of C. knabi for PG-rich willow plants because larvae grew and survived better and that selection by common exotic generalist predators would reinforce this preference.     

Do native predators benefit from non‐native prey?

Despite knowledge on invasive species’ predatory effects, we know little of their influence as prey. Non‐native prey should have a neutral to positive effect on native predators by supplementing the prey base. However, if non‐native prey displace native prey, then an invader's net influence should depend on both its abundance and value relative to native prey. We conducted a meta‐analysis to quantify the effect of non‐native prey on native predator populations. Relative to native prey, non‐native prey similarly or negatively affect native predators, but only when studies employed a substitutive design that examined the effects of each prey species in isolation from other prey. When native predators had access to non‐native and native prey simultaneously, predator abundance increased significantly relative to pre‐invasion abundance. Although non‐native prey may have a lower per capita value than native prey, they seem to benefit native predators by serving as a supplemental prey resource.     

Recognition and Response to Native and Novel Predators in the Largespring mosquitofish,Gambusia geiseri

The introduction of predator species into new habitats is an increasingly common consequence of human activities, and the persistence of native prey species depends upon their response to these novel predators. In this study, we examined whether the Largespring mosquitofish, Gambusia geiseri exhibited antipredator behavior and/or an elevation of circulating stress hormones (cortisol) to visual and chemical cues from a native predator, a novel predator, or a non‐predatory control fish. Prey showed the most pronounced antipredator response to the native predator treatment, by moving away from the stimulus, while the prey showed no significant changes in their vertical or horizontal position in response to the novel or non‐predator treatments. We also found no significant difference in water‐borne cortisol release rates following any of the treatments. Our results suggest the prey did not recognize and exhibit antipredator behavior to the novel predator, and we infer that this predator species could be detrimental if it expands into the range of this prey species. Further, our study demonstrates prey may not respond to an invasive predator that is phylogenetically, behaviorally, and morphologically dissimilar from the prey species' native predators.     

Naïveté in novel ecological interactions: lessons from theory and experimental evidence

The invasion of alien species into areas beyond their native ranges is having profound effects on ecosystems around the world. In particular, novel alien predators are causing rapid extinctions or declines in many native prey species, and these impacts are generally attributed to ecological naïveté or the failure to recognise a novel enemy and respond appropriately due to a lack of experience. Despite a large body of research concerning the recognition of alien predation risk by native prey, the literature lacks an extensive review of naïveté theory that specifically asks how naïveté between novel pairings of alien predators and native prey disrupts our classical understanding of predator–prey ecological theory. Here we critically review both classic and current theory relating to predator–prey interactions between both predators and prey with shared evolutionary histories, and those that are ecologically ‘mismatched’ through the outcomes of biological invasions. The review is structured around the multiple levels of naïveté framework of Banks & Dickman (2007), and concepts and examples are discussed as they relate to each stage in the process from failure to recognise a novel predator (Level 1 naïveté), through to appropriate (Level 2) and effective (Level 3) antipredator responses. We discuss the relative contributions of recognition, cue types and the implied risk of cues used by novel alien and familiar native predators, to the probability that prey will recognise a novel predator. We then cover the antipredator response types available to prey and the factors that predict whether these responses will be appropriate or effective against novel alien and familiar native predators. In general, the level of naïveté of native prey can be predicted by the degree of novelty (in terms of appearance, behaviour or habitat use) of the alien predator compared to native predators with which prey are experienced. Appearance in this sense includes cue types, spatial distribution and implied risk of cues, whilst behaviour and habitat use include hunting modes and the habitat domain of the predator. Finally, we discuss whether the antipredator response can occur without recognition per se, for example in the case of morphological defences, and then consider a potential extension of the multiple levels of naïveté framework. The review concludes with recommendations for the design and execution of naïveté experiments incorporating the key concepts and issues covered here. This review aims to critique and combine classic ideas about predator–prey interactions with current naïveté theory, to further develop the multiple levels of naïveté framework, and to suggest the most fruitful avenues for future research.     

Eating their way to the top? Mechanisms underlying the success of invasive insect generalist predators

Insect generalist predators have been introduced outside of their native range intentionally to improve biological control, or accidentally during commerce, and can subsequently become invasive. Invasive insect generalist predators (IIGP) have widespread impacts on invaded communities because they consume both herbivores and other predators. Also, they often reach higher densities than and displace similar native species. Reflecting the complexity of their ecological roles, a wide variety of mechanisms might contribute to invasive success by IIGP. These species often drive resources to lower levels than do natives, leading to intense resource competition and sometimes competitive exclusion of other predators. The broader range of resources used by many IIGP can heighten their competitive advantage, particularly when IIGP exploit modified habitats. In either case, IIGP improve herbivore suppression by depressing prey densities below pre-invasion levels. Coexistence among native and invasive generalists is fostered when species differ substantially in their niche requirements. In this case, a larger proportion of the total prey population is subject to attack post- than pre-invasion, which strengthens prey suppression. On the other hand, some IIGP feed heavily on other predators, at times leading to a weakening of prey suppression. Future research should continue to explore the roles of competition and niche partitioning on larger spatial scales, and in both the native and invasive ranges of IIGP. Additionally, combining data from empirical studies with theory might be an effective way to predict the spread and community impacts of IIGP invasions.     

Damage, digestion, and defence: the roles of alarm cues and kairomones for inducing prey defences

Inducible defences are widely used for studying phenotypic plasticity, yet frequently we know little about the cues that induce these defences. For aquatic prey, defences are induced by chemical cues from predators (kairomones) and injured prey (alarm cues). Rarely has anyone determined the separate and combined effects of these cues, particularly across phylogenetically diverse prey types. We examined how tadpoles (Hyla versicolor) altered their defences when 10 different prey were either crushed by hand or consumed by predators. Across all prey types, crushing induced only a subset of the defences induced by consumption. Consuming vs. crushing produced additive responses for behaviour but synergistic responses for morphology and growth. Moreover, we discovered the first extensive evidence that prey responses to different alarm cues depends on prey phylogeny. These results suggest that the amount of information available to the prey affects both the quantitative and qualitative nature of the defended phenotype.