Consequences of infanticide for a gregarious ectoparasitoid of leafroller larvae

1. In this laboratory study, the clutch size and handling time of Goniozus jacintae were investigated, a comparison of its life‐history performance between primary and secondary (laid after infanticide events) broods was carried out, and the lipid and protein concentrations in the haemolymph of non‐parasitised and parasitised hosts were estimated. 2. It was found that G. jacintae temporarily paralysed its host larvae for 66 min and briefly guarded its brood for 66 min. The clutch size of G. jacintae increased from two to seven with increasing larval fresh weight of its host, and both ovicide and larvicide of primary clutches occurred in 81% of encounters. 3. Secondary clutches of G. jacintae were significantly larger than primary clutches in two of three ovicide treatments for the same host individuals. Secondary clutches also experienced greater brood survivorship than primary clutches. 4. Lipid concentrations were consistently higher in the haemolymph of parasitised hosts, and protein concentrations were initially higher (until egg hatch), but increased at a lower rate in parasitised hosts than in non‐parasitised hosts. 5. This study is the first to provide evidence that improved nutritional quality could be an important benefit of infanticide for an insect parasitoid, allowing for larger clutch size and improved brood survivorship among secondary broods.     

To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti‐parasite strategies

Hosts either tolerate avian brood parasitism or reject it by ejecting parasitic eggs, as seen in most rejecter hosts of common cuckoos, Cuculus canorus, or by abandoning parasitized clutches, as seen in most rejecter hosts of brown‐headed cowbirds, Molothrus ater. What explains consistent variation between alternative rejection behaviours of hosts within the same species and across species when exposed to different types of parasites? Life history theory predicts that when parasites decrease the fitness of host offspring, but not the future reproductive success of host adults, optimal clutch size should decrease. Consistent with this prediction, evolutionarily old cowbird hosts, but not cuckoo hosts, have lower clutch sizes than related rarely‐ or newly parasitized species. We constructed a mathematical model to calculate the fitness payoffs of egg ejector vs. nest abandoner hosts to determine if various aspects of host life history traits and brood parasites’ virulence on adult and young host fitness differentially influence the payoffs of alternative host defences. These calculations showed that in general egg ejection was a superior anti‐parasite strategy to nest abandonment. Yet, increasing parasitism rates and increasing fitness values of hosts’ eggs in both currently parasitized and future replacement nests led to switch points in fitness payoffs in favour of nest abandonment. Nonetheless, nest abandonment became selectively more favourable only at lower clutch sizes and only when hosts faced parasitism by a cowbird‐ rather than a cuckoo‐type brood parasite. We suggest that, in addition to evolutionary lag and gape‐size limitation, our estimated fitness differences based on life history trait variation provide new insights for the consistent differences observed in the anti‐parasite rejection strategies between many cuckoo‐ and cowbird‐hosts.     

Influence of host size on oviposition behaviour and fitness of Elachertus cacoeciae attacking a low-density population of spruce budworm Choristoneura fumiferana larvae

1. Oviposition behaviour and host size ? fitness relationships of a gregarious, idiobiont ectoparasitoid, Elachertus cacoeciae (Howard) (Hymenoptera: Eulophidae), were studied by implanting one fourth‐, fifth‐, and sixth‐instar spruce budworm Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) larva per tree in a stand where the density of the wild C. fumiferana population was low. 2. Development time of E. cacoeciae larvae was quickest on fifth‐instar C. fumiferana larvae, which were the preferred hosts for oviposition. 3. Brood sex ratio (proportion of females) was related positively to increasing C. fumiferana instar, indicating that more females were laid on larger hosts. 4. Parasitoid offspring size increased with increasing C. fumiferana instar and decreased with increasing brood size on smaller hosts. Female but not male size was related positively to increasing brood sex ratio (proportion of females). 5. Under laboratory conditions, parasitoid longevity was related positively to parasitoid size and realised lifetime fecundity, and clutch size was related positively to host size. 6. These results suggest that selection of intermediate‐sized C. fumiferana larvae may be adaptive for E. cacoeciae.     

Costs and consequences of superparasitism in the polyembryonic parasitoid Copidosoma koehleri (Hymenoptera: Encyrtidae)

Abstract.  1. Polyembryonic wasps provide dramatic examples of intra-specific developmental conflict. In these parasitoids, each egg proliferates into a clonal lineage of genetically identical larvae. If more than one egg is laid in a host (superparasitism), individuals of different clones may compete for food resources.
2. In the polyembryonic encyrtid Copidosoma koehleri , one larva per clone can differentiate into a sterile soldier. It is shown that soldiers are always females, and that they attack intra-specific competitors.
3. Research hypotheses were that (a) clones that develop in superparasitised hosts suffer heavier mortality than clones that develop in singly parasitised hosts, and (b) female clones cause higher mortality to their competitors than male clones, hence larval survival is lower in superparasitised hosts that contain females than in male-only broods.
4. The potential frequency of superparasitism in C. koehleri was manipulated by varying parasitoid–host ratios and exposure durations.
5. As parasitoid densities and exposure durations increased, the frequency of superparasitism rose, brood sizes increased, but the number of hosts that completed development was reduced. The number of offspring per parasitoid female decreased with increasing parasitoid–host ratios. Offspring size and longevity were inversely correlated with brood size. As superparasitism rates increased, fewer all-male broods were produced. Male–female broods were female-biased, suggesting selective killing of males by female soldiers. All-female broods were significantly smaller than all-male broods at high parasitoid densities only, possibly reflecting aggression among soldiers of competing clones.
6. The results support the working hypotheses, and suggest that female larvae outcompete males in superparasitised hosts.     

The developmental strategies and related profitability of an idiobiont ectoparasitoid Sclerodermus pupariae vary with host size

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Host location by the fruit fly parasitoid Diachasmimorpha krausii: role of fruit fly species,life stage and host plant

• 1 Diachasmimorpha krausii is a braconid parasitoid of larval tephritid fruit flies, which feed cryptically within host fruit. At the ovipositor probing stage, the wasp cannot discriminate between hosts that are physiologically suitable or unsuitable for offspring development and must use other cues to locate suitable hosts.
• 2 To identify the cues used by the parasitoid to find suitable hosts, we offered, to free flying wasps, different combinations of three fruit fly species (Bactrocera tryoni, Bactrocera cacuminata, Bactrocera cucumis), different life stages of those flies (adults and larvae) and different host plants (Solanum lycopersicon, Solanum mauritianum, Cucurbita pepo). In the laboratory, the wasp will readily oviposit into larvae of all three flies but successfully develops only in B. tryoni. Bactrocera tryoni commonly infests S. lycopersicon (tomato), rarely S. mauritianum (wild tobacco) but never C. pepo (zucchini). The latter two plant species are common hosts for B. cacuminata and B. cucumis, respectively.
• 3 The parasitoid showed little or no response to uninfested plants of any of the test species. The presence of adult B. tryoni, however, increased parasitoid residency time on uninfested tomato.
• 4 When the three fruit types were all infested with larvae, parasitoid response was strongest to tomato, regardless of whether the larvae were physiologically suitable or unsuitable for offspring development. By contrast, zucchini was rarely visited by the wasp, even when infested with B. tryoni larvae.
• 5 Wild tobacco was infrequently visited when infested with B. cacuminata larvae but was more frequently visited, with greater parasitoid residency time and probing, when adult flies (either B. cacuminata or B. tryoni) were also present.
• 6 We conclude that herbivore‐induced, nonspecific host fruit wound volatiles were the major cue used by foraging D. krausii. Although positive orientation to infested host plants is well known from previous studies on opiine braconids, the failure of the wasp to orientate to some plants even when infested with physiologically suitable larvae, and the secondary role played by adult fruit flies in wasp host searching, are newly‐identified mechanisms that may aid parasitoid host location in environments where both physiologically suitable and unsuitable hosts occur.

     

Host influences on sex ratio, longevity, and egg load of two Metaphycus species parasitic on soft scales: implications for insectary rearing

Metaphycus flavus (Howard) and M. stanleyi Compere (Hymenoptera: Encyrtidae) are currently being screened for use as augmentative biological control agents of citrus-infesting soft scales (Homoptera: Coccidae). Two factors were investigated, host quality-dependent sex allocation and local mate competition, which likely influence these parasitoid's sex allocation strategies and are therefore of interest for their mass-rearing. The results of these studies suggested that, under the mass-rearing protocol that is envisioned for these parasitoids, offspring sex ratios in both M. flavus and M. stanleyi are dominated by host quality (= size) influences, but not by interactions with other females. These results indicated that host size strongly influences offspring sex ratios and brood sizes; larger hosts led to more female offspring and larger broods. In contrast, increasing the number of parental females did not lead to fewer female offspring as expected under local mate competition. Additionally, within-brood sex ratios did not vary with brood size; this result is inconsistent with expected sex ratios due to local mate competition. Other results also indicated that host quality was a dominant influence on M. flavus' and M. stanleyi's sex ratios. Larger hosts led to a larger size in the emerging wasps, and larger wasps had greater egg loads and lived longer than smaller wasps. However, wasp longevity, and the influence of wasp size on longevity were mediated by a wasp's diet. Metaphycus flavus females lived the longest when they had access to hosts, honey, and water, followed by honey and water, and shortest when they had access to water alone; M. stanleyi females lived longest with honey and water, followed by hosts, honey, and water, and shortest with water alone. Greater wasp size led to greater longevity in females only when they had access to food (honey, or hosts and honey). Finally, other results suggested that both M. flavus and M. stanleyi are facultatively gregarious. Wasp size did not decrease with brood size as expected under superparasitism. Overall, the results of these studies suggested that holding newly emerged females of both M. flavus and M. stanleyi for several days in the presence of an appropriate food source before field release could enhance a female's performance as an augmentative biological control agent. It increases their initial life expectancy following release, and maximizes the females' egg load (both Metaphycus species) and resources for replacing oviposited eggs (M. flavus only).     

Host population structure affects field sex ratios of the heteronomous hyperparasitoid, Coccophagus atratus

Abstract.

• 1 Second instar Filippia gemina de Lotto scale insects are the preferred hosts of female Coccophugus atrutus Compere larvae. These scale insects were found on their host plants, Chrysanthemoides monilifera Norlindh and Cliffortia strobilifera Mettenius, only at certain times during a 1 year sampling programme.
• 2 Late larval instars and prepupae of C.atratus, and a Metaphycus species, are the preferred hosts of male C.atratus larvae. These hosts, although they occurred on the same host plants as hosts for female C.atratus, were most numerous at different times during the sampling period.
• 3 The ratio of hosts suitable for C.atratus varied from a predominance of hosts suitable for females through to a predominance of hosts suitable for males. Sex ratios of adult C.atratus followed a similar trend but did not reflect, exactly, the ratio of available hosts. Differences in mortality between sexes and hyperparasitism may account for this anomaly.
• 4 Variable population sex ratios observed in C.atratus apparently result from the behaviour of individual females in which brood sex ratios are dependent on the relative availability of hosts for males and hosts for females. This behaviour, in turn, may result from variability in the host population structure but may also result from selection pressures operating at the time that heteronomous hyperparasitism evolved.

     

Sequences of sex allocation and mortality in clutches of Metaphycus parasitoids of soft scale insects and the prevalence of all‐female broods

1. In many gregarious or quasi‐gregarious parasitoids that experience local mate competition, precise sex ratios with low variance are observed. Precise sex ratios can be achieved by laying male and female eggs in non‐random sequences. 2. Developmental mortality can also alter sex ratios of emerging offspring, and subsequently influence sex ratio optima. 3. The present study investigates sex allocation by Metaphycus flavus Howard, M. luteolus Timberlake, and M. angustifrons Compere (Hymenoptera: Encyrtidae), endoparasitoids of soft scale insects, in the laboratory. 4. All three Metaphycus species had precise secondary sex ratios when parasitising brown soft scale, Coccus hesperidum, L. in the laboratory. Moreover, we documented that all three species lay fertilised (= female) eggs first followed by unfertilised (= male) eggs at the end of the oviposition bout. However, there were significant differences in sex allocation sequences among species. 5. Mortality rates of eggs allocated within an oviposition bout also varied considerably, indicating that there is a significant interspecific variation in sequence position‐specific mortality. 6. Using a stochastic Monte Carlo simulation approach, we provide evidence that the incidence of all‐female broods in these parasitoid wasps appears mainly due to developmental mortality and not due to decisions by the ovipositing female. In two species the prevalence of all‐female broods was independent of clutch size, contrary to what is expected from theory. The influence of mortality on optimal sex allocation in these parasitoids is discussed.     

Great Spotted Cuckoos Frequently Lay Their Eggs While Their Magpie Host is Incubating

Species that suffer from brood parasitism face a considerable reduction in their fitness which selects for the evolution of host defences. To prevent parasitism, hosts can mob or attack brood parasites when they approach the host nest and block the access to the nest by sitting on the clutch. In turn, as a counter‐adaptation, brood parasites evolved secretive behaviours near their host nests. Here, we have studied great spotted cuckoo (Clamator glandarius) egg‐laying behaviour and defence by their magpie (Pica pica) hosts inside the nest using continuous video recordings. We have found several surprising results that contradict some general assumptions. The most important is that most (71%) of the parasitic events by cuckoo females are completed while the magpie females are incubating. By staying in the nest, magpies force cuckoo females to lay their egg facing the high risk of being attacked by the incubating magpie (attack occurred in all but one of the events, n = 15). During these attacks, magpies pecked the cuckoo violently, but could never effectively avoid parasitism. These novel observations expand the sequence of adaptations and counter‐adaptations in the arms race between brood parasites and their hosts during the pre‐laying and laying periods.     

An asexual strain of Lysiphlebus fabarum (Hymenoptera: Braconidae) gains fitness in superparasitised hosts,but a sexual strain pays costs

1. The fitness impacts of two levels of superparasitism were compared in a sexual and an asexual strain of Lysiphlebus fabarum (Marshall) developing in black bean aphids. 2. Asexual females were larger than sexuals, had higher eggs loads, and better adult emergence in two of three treatments, but parasitised fewer available aphids in 24 h (80% fewer when foraging alone). 3. Superparasitism resulted in wasps that were larger than those emerging from singly parasitised aphids, and asexual females had larger ovaries, without delayed development. In contrast, sexual wasps took about 2 days longer to develop in superparasitised hosts, and females had smaller ovaries with lower egg loads. 4. The growth of host aphids bearing asexual larvae was not reduced relative to healthy aphids, except when heavily superparasitised, whereas sexual larvae reduced aphid growth in all treatments. 5. Elimination of supernumerary larvae was virtually complete at 4.5 days after parasitism by sexual females, but no elimination occurred among asexual larvae. The evolution of diminished aggression in asexual larvae may be facilitated by greater genetic similarity, without necessarily leading to gregarious development, assuming the extended life of supernumeraries somehow improves survivor fitness. 6. Sons of virgin sexual females developed faster than sons of mated females, implicating parental effects that slowed development of the latter. 7. It is concluded that asexual females of this strain use superparasitism to improve host quality, and thus the fitness of their daughters, whereas females of the sexual strain prioritise offspring number.     

Reproductive and subsocial behaviour in the ovoviviparous leaf beetle Gonioctena sibirica (Coleoptera: Chrysomelidae)

Abstract.

• 1 In the spring, females of the leaf beetle Gonioctena sibirica deposited larvae on the ventral surface of growing young leaves situated on the apical position of shoots of the willow Salix bakko.
• 2 The parent females remained with the larvae usually on the underside of the basal part of leaves, facing toward the base of shoots. When other arthropods approached, the females temporarily moved towards these intruders, showing aggressive behaviour such as swinging the body or stamping the legs. Many females remained with their larvae until the larvae grew into the final (fourth) instar. No female produced an additional brood in the field.
• 3 Broods from which parent females were experimentally removed suffered higher mortality than those in which females were left intact. Arthropods such as spiders and ants were observed preying on the larvae. In contrast, the survivorship of broods from which females were removed and intruders were excluded with a sticky substance applied to the base of twigs was not different from that of control broods. These results demonstrate that the main mortality factor of offspring is pedestrian arthropod predators and females physically repel the predators.
• 4 Potentially alternative reproductive strategies, such as producing a large number of offspring by iteroparity and/or larger brood(s) with less or no care, seem to be inhibited in G.sibirica by larval dependence on growing young leaves which are temporally limited and by ovoviviparity which may have limited brood size.

     

Adaptive decision making or differential mortality: what causes offspring emergence in a gregarious parasitoid?

Hosts represent a limited resource for the developing offspring of parasitic insects laying eggs in or on spatially discrete resources like fruits, seeds, or other insects. The quality of hosts differs with respect to the value and amount of resources they provide for the feeding larvae. Accordingly, the size of a clutch of eggs laid on a given host should be a function of host quality, because severe competition between developing larvae can lead to increased mortality and/or decreased size of the offspring, both causing a fitness loss for the offspring and the mother. Therefore, females should be selected for the ability to estimate host quality and to adjust their clutch size accordingly. Using the parasitic wasp Nasonia vitripennis (Walker) (Hymenoptera: Pteromalidae) this study investigated the respective contribution of developmental mortality of offspring vs. the clutch size decision of the mother as a determinant of final offspring emergence per host. In addition, taking offspring size into account, the study examined the fitness consequences of female oviposition decisions. Developmental mortality was very low in all quality classes of hosts except previously frozen and thus dead host pupae. Females laid reduced clutch sizes on dead, previously parasitized, and smaller hosts. In contrast to offspring number, offspring size did not differ between host qualities. We conclude that females are able to sense the quality of a host and adjust the number of eggs they lay to mitigate larval competition.     

MAGPIE HOST MANIPULATION BY GREAT SPOTTED CUCKOOS: EVIDENCE FOR AN AVIAN MAFIA?

Why should the hosts of brood parasites accept and raise parasitic offspring that differ dramatically in appearance from their own? There are two solutions to this evolutionary enigma. (1) Hosts may not yet have evolved the capability to discriminate against the parasite, or (2) parasite-host systems have reached an evolutionary equilibrium. Avian brood parasites may either gain renesting opportunities or force their hosts to raise parasitic offspring by destroying or preying upon host eggs or nestlings following host ejection of parasite offspring. These hypotheses may explain why hosts do not remove parasite offspring because only then will hosts avoid clutch destruction by the cuckoo. Here we show experimentally that if the egg of the parasitic great spotted cuckoo Clamator glandarius is removed from nests of its magpie Pica pica host, nests suffer significantly higher predation rates than control nests in which parasite eggs have not been removed. Using plasticine model eggs resembling those of magpies and observations of parasites, we also confirm that great spotted cuckoos that have laid an ejected egg are indeed responsible for destruction of magpie nests with experimentally ejected parasite eggs. Cuckoos benefit from destroying host offspring because they thereby induce some magpies to renest and subsequently accept a cuckoo egg.     

Clutch size in frugivorous insects as a function of host firmness: the case of the tephritid fly Anastrepha ludens

Abstract. 1. Optimal clutch size theory predicts that individuals will oviposit the number of eggs that increases their fitness. In Anastrepha ludens Loew (Diptera: Tephritidae), females oviposit larger clutches in unripe (firm) fruits than in ripe (soft) fruits. The following hypotheses were tested: (1) Using fruit firmness as an indicator of fruit quality, A. ludens females vary the number of eggs per clutch every time they reach an oviposition decision. (2) Maximising offspring survival with respect to either unripe or ripe fruit requires placing large clutches in firm fruit and smaller clutches in soft fruit. 2. Agar spheres were used as artificial hosts. Three agar concentrations resulted in three degrees of firmness. Mango fruits Mangifera indica L. served as natural hosts. Ripe and unripe fruits were used to test soft and firm host conditions respectively. Females laid significantly larger clutches in the firmer artificial hosts than in the softer hosts. They also laid significantly more eggs in artificial hosts without sugar than in hosts with sugar. Firm (unripe) mangoes also received significantly larger clutches than soft (ripe) mangoes. 3. When an individual female was first presented with a firm artificial host, it laid a large clutch. If subsequently offered a soft host, the female laid a significantly smaller clutch. Finally, if again offered a firm host, clutch size was increased significantly. 4. Possible trade‐offs in offspring fitness were explored in ripe and unripe mangoes by measuring offspring egg‐to‐adult survival, pupal weight, mean adult longevity, and fecundity. Despite the fact that larval survival was greater in soft fruit than in firm fruit, parameters such as pupal weight, mean longevity, and fecundity of adults stemming from both fruit types did not differ significantly. 5. A probable trade‐off between high offspring mortality caused by host unsuitability and low offspring and adult mortality caused by parasitism and predation is discussed as the reason for the exploitation of sub‐optimal hosts.     

Brood guarding in a bethylid wasp

Abstract.

• 1 Atypical of the parasitoid Hymenoptera, female Goniozus nephantidis Muesebeck remain with offspring until they pupate. This behaviour will only have evolved if the consequent reduction in fecundity is outweighed by fitness returns.
• 2 G.nephantidis competes for hosts with conspecific and non-conspecific parasitoids. The effectiveness of G.nephantidis at deterring superparasitism and multiparasitism is tested. Brood survivorships were compared when G.nephantidis and Bracon hebetor Say intrude on hosts with the mother present and absent and with offspring at different developmental stages.
• 3 Laying by the intruder reduced brood survivorship. Guarding reduced oviposition on unparasitized hosts by intruding females, and prevented superparasitism of hosts with egg broods. Hosts with larval broods were rarely superparasitized. Multiparasitism was common except on hosts with guarded larval broods, but even here survivorship was reduced.
• 4 Competitive asymmetries determined the outcome of contests for possession of host resources.
• 5 The costs of remaining seem at least partially offset by the prevention of oviposition by competing parasitoids.

     

Repeatability of Host Female and Male Aggression Towards a Brood Parasite

Current research on behavioural consistency showed that various types of animal behaviour are highly repeatable in the context of mate choice, exploration and parental care, including nest protection. However, the repeatability of aggressive nest defence has not yet been studied in hosts of brood parasites, although host aggression against adult parasites represents a crucial line of antiparasitic defences. Here, we investigated the between‐season repeatability of the great reed warbler (Acrocephalus arundinaceus) aggression towards a stuffed dummy of the brood parasitic common cuckoo (Cuculus canorus). We found that under the relatively stable risk of brood parasitism across breeding seasons, female responses to the cuckoo were highly repeatable, whereas male responses were variable. We suggest that the potential explanation for the observed patterns of female and male behaviours may lie in female's prominent roles in offspring care and nest protection, and in her lower renesting potential in comparison with that of males. However, further studies on the relationship between host aggression and other types of behaviours (host personality) and their fitness consequences under the fluctuating parasitism pressures are required to clarify the adaptive significance of aggressive behaviour observed in hosts of brood parasites.     

RETRACTED ARTICLE: Foraging of Host-Habitat and Superparasitism in <Emphasis Type="Italic">Cotesia glomerata</Emphasis>: A Gregarious Parasitoid of <Emphasis Type="Italic">Pieris brassicae</Emphasis>

The study investigates differences in the oviposition pattern of a braconid parasitoid, Cotesia glomerata (Linn.) in Pieris brassicae (Linn.) in relation to their use of different cruciferous food plants. The response of P. brassicae to superparasitism and consequences for the parasitoid were examined in order to elucidate the ecological significance of this behaviour. Female parasitoid located various crucifers and searched for host more frequently almost on all the host plants tested i.e. cabbage, cauliflower, Chinese cabbage, broccoli and radish. According to the estimated relative number of female locating hosts, cabbage was the most attractive plant for C. glomerata and total number of eggs laid in host larvae feeding on it was higher than in larvae feeding on other plants. Laboratory experiments demonstrated that superparasitism reduced survivorship of P. brassicae larvae. Superparasitism lengthened parasitoid development and prolonged the feeding period of host larvae. Sex ratio and the body weight of emergent wasps correlated negatively with brood size. Despite a trade-off between maximising brood size and optimising the fitness of individual offspring, two or three ovipositions on P. brassicae larvae resulted in a greater female dry mass than did a single oviposition on the host. Thus, superparasitism might be of adaptive significance under certain circumstances, especially when host density is low and unparasitized hosts are rare in a habitat.     

Host activity and wasp experience affect parasitoid wasp foraging behaviour and oviposition on nematode‐infected larvae of the forestry pest Hylobius abietis

1. Entomopathogenic nematodes (EPN) are currently being used as introduced biological control agents against the larvae of the native European forestry pest Hylobius abietis L. which develop under the bark of stumps and roots of newly dead conifer trees. 2. The potential for resource competition between gregarious ectoparasitoid Bracon hylobii Ratz and EPN by recording oviposition and related behaviours of B. hylobii females on EPN‐infected H. abietis larvae was investigated. Wasps did not parasitise EPN‐infected host larvae that were dead when presented, but naÏve and experienced wasps parasitised live EPN‐infected hosts. NaÏve wasps parasitised live EPN‐infected hosts significantly less frequently than healthy hosts only when the infected larvae were close to death (i.e. died during 24‐h trial). Parasitism by experienced wasps was unaffected by host infection. 3. Wasp probing and oviposition were positively associated with the amount of host movement. Preventing H. abietis larvae from chewing on bark significantly reduced parasitism by naÏve, but not experienced wasps. 4. The number of eggs per clutch was not affected by bark chewing or EPN‐infection of H. abietis larvae. 5. NaÏve and experienced B. hylobii parasitised two abnormal hosts (larvae of coleopteran Rhagium bifasciatum Fabricius and lepidopteran Galleria mellonella L.), both of which moved and chewed on bark during trials. 6. It was concluded that B. hylobii can use vibrational cues generated by host movement and feeding to locate hosts at short range and accepts unsuitable (EPN‐infected or abnormal) hosts as long as these create such cues. The implications for competition between B. hylobii and EPN and possible ways of minimising it when applying EPN are discussed.     

Competitor avoidance drives within‐host feeding site selection in a passively dispersed herbivore

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