WASP‐ASSOCIATED FACTORS ACT IN INTERSPECIES COMPETITION DURING MULTIPARASITISM

Coexistence or displacement of parasitoids in hosts during intrinsic competitive interactions between different parasitoid species (multiparasitism) may depend on their life history traits and behavior. Intense competition for possession of hosts may lead to the elimination of the inferior competitor through physical attack and/or physiological suppression. However, the mechanisms of physiological suppression during multiparasitism remain unclear. Previous work has shown that first instar larvae of the solitary endoparasitoid Meteorus pulchricornis possess well‐developed mandibles that are used to kill competitors. Two gregarious endoparasitoids, Cotesia kariyai and C. rufricus, share host resources especially when the time gap of oviposition is short. Here, we investigated the physiological influence of wasp‐regulatory factors of the three endoparasitoids, M. pulchricornis, C. kariyai, and C. ruficrus, in their common host Mythimna separata. We found that MpVLP alone (or with venom) deleteriously affected the development of the two gregarious species. Similarly, CkPDV plus venom had toxic effect on M. pulchricornis eggs and immature larvae, although they were not harmful to immature stages of C. ruficrus. Cotesia kariyai and C. ruficrus were able to coexist mainly through the expression of regulatory factors and both could successfully emerge from a multiparasitized host. The injection of CkPDV plus venom after oviposition in L5 host larvae facilitated C. ruficrus development and increased the rate of successful parasitism from 9% to 62%. This suggests that the two gregarious parasitoid wasps exhibit strong phylogenetic affinity, favoring their coexistence and success in multiparasitized hosts.     

Cover Caption

In biological control of Paratrioza sinica, nymphs are important stages because of both ecto‐ and endoparasitoid which act as effective agents. However, endoparasitoid is often trapped fatally by P. sinica nymphs due to strong host immunity. The cover photo shows P. sinica nymphs on the Lycium barbarum leave surface and endoparasitoids act inside the body of nymphs. Around them, some egg shells with stalks of P. sinica are standing on the wolfberry leave (see pages 815‐825). Photo provided Run‐Zhi Zhang.     

Responses of potential hosts of Asian cuckoos to experimental parasitism

In the arms race between avian brood parasites and their hosts, several adaptations and counter‐adaptations have evolved. The most prominent host defence is rejection of parasitic eggs. We experimentally parasitized nests of 10 potential host species breeding in sympatry with four different cuckoo species in an area in Bangladesh using differently coloured model eggs to test host responses. In four species we introduced both mimetic and non‐mimetic eggs. Black Drongos Dicrurus macrocercus, hosts of the Indian Cuckoo Cuculus micropterus, rejected all model eggs. Common Mynas Acridotheres tristis and Jungle Babblers Turdoides striata accepted all eggs regardless of mimicry. These two species are parasitized by Asian Koels Eudynamys scolopaceus, Common Hawk‐cuckoo Hierococcyx varius and, in the case of Jungle Babblers, Jacobin Cuckoos Clamator jacobinus. Pied Mynas Gracupica contra, with no records of parasitism in our study area, also accepted all eggs regardless of mimicry. In the six remaining species, all of which lay spotted eggs, we introduced only non‐mimetic eggs. Black‐hooded Orioles Oriolus xanthornus rejected all model eggs, even though we have found no records of natural parasitism. Long‐tailed Shrikes Lanius schach and House Crows Corvus splendens, hosts of Asian Koels, rejected 75 and 9.1% of model eggs, respectively. Large‐billed Crows Corvus macrorhynchos, apparently not used as hosts in our study area, accepted all blue but rejected all brown model eggs. Oriental Magpie‐Robins Copsychus saularis and Red‐vented Bulbuls Pycnonotus cafer accepted all non‐mimetic model eggs. In Black Drongos, Long‐tailed Shrikes and Black‐hooded Orioles, all model eggs were ejected within 24 h of introduction. The results show considerable variation in egg rejection rates among various species, providing baseline data for further investigation of co‐evolutionary interactions between brood parasites and hosts in this region.     

Multiparasitism of Choristoneura fumiferana by the ichneumonid Tranosema rostrale and the tachinid Actia interrupta: occurrence in the field and outcome of competition under laboratory conditions

Tranosema rostrale (Brishke) (Hymenoptera, Ichneumonidae) and Actia interrupta Curran (Hymenoptera: Tachinidae) are the two endoparasitoids most frequently encountered in low-density populations of the spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera, Tortricidae), in the Quebec City region. Monitoring of attack rates of implanted C. fumiferana larvae at two different study sites suggested the possible existence of competition between the two parasitoids, with A. interrupta seemingly displacing T. rostrale. Here, we show that multiparasitism involving these two species does occur in the field, but at a frequency too low to explain the seasonal pattern of decline in apparent parasitism by T. rostrale that accompanies the rise of A. interrupta attack rates. We also provide preliminary evidence, from laboratory experiments, that A. interrupta has a competitive advantage over T. rostrale and that the success of parasitism by A. interrupta may be enhanced by prior parasitism by T. rostrale under certain conditions, possibly due to the presence of the latter species' polydnavirus. In addition, we describe a PCR-based method that we developed to help detect the presence of T. rostrale eggs which often escape detection by simple visual examination of the dissected host larvae; DNA sequences specific to the polydnavirus injected by the female wasp at the time of oviposition can be readily amplified from whole host larvae.     

Interspecific competition between two endoparasitoids Cotesia vestalis (Hymenoptera: Braconidae) and Oomyzus sokolowskii (Hymenoptera: Eulophidae)

Two endoparasitoids, Cotesia vestalis and Oomyzus sokolowskii, parasitize the same host, larvae of Plutella xylostella. These two species have evolved different parasitization strategies. O. sokolowskii expresses a single factor, venom, and exerts virtually no detrimental effects on the development of its host. C. vestalis, on the other hand, injects polydnavirus (PDV) and venom during oviposition, and teratocytes are released into the host's hemolymph after egg hatching. Parasitization suppresses host immune reactions and redirects its developmental program. Because both these species parasitize the same stage of their hosts, there is the possibility of multiparasitism in nature. Only one species survives multiparasitism and because of its parasitic strategy, we hypothesized that C. vestalis would invariably be the stronger competitor. We designed competition experiments which revealed that C. vestalis is a stronger competitor than O. sokolowskii. We also show that C. vestalis survives intrinsic competition with O. sokolowskii through two mechanisms: physical attack and physiological suppression. We discovered melanized wounds on O. sokolowskii eggs and larvae, which is strong evidence of physical attacks. The physiological suppression is due to PDV and venom injected by C. vestalis. To test this idea more rigorously, we designed a pseudoparasitization experiment which revealed that no O. sokolowskii emerged from multiparasitized hosts when infertile C. vestlais eggs and normal O. sokolowskii larvae are both present inside the same host. These results support our hypothesis that C. vestalis is the stronger competitor and demonstrate two mechanisms that account for the outcome of intrinsic competition between these two endoparasitoids.     

Techniques for rearing tachinid parasitoids of the European Earwig Forficula auricularia

Two European species of the family Tachinidae (Diptera), Triarthria setipennis and Ocytata pallipes, whose larvae live as endoparasitoids of the European earwig (Forficula auricularia), were studied to develop experimental methods to improve laboratory production of the parasitoids. More than 84% parasitism was achieved by inoculating with larvae of T. setipennis when earwigs were immobilized. Lower rates of parasitism were obtained when earwigs were mobile and could fend off attacking parasitoid larvae before they could penetrate the host. For O. pallipes, where the microtype eggs have to be eaten by the host, incubation of starved earwigs with food items carrying 3–5 parasitoid eggs resulted in 60% parasitism.     

Interspecific Relations Between Two Sympatric Species of Hymenoptera, Dinarmus basalis (Rond) and Eupelmus vuilleti (Crw), Ectoparasitoids of the Bruchid Callosobruchus maculatus (F)

Dinarmus basalis (Rond) and Epelmus vuilleti (Crw) are two Hymenopteran species, which are solitary ectoparasitoids of bruchid larvae. In the presence of seeds of Vigna unguiculata (Walp) containing hosts parasitized by E. vuilleti, a high percentage of D. basalis females avoided multiparasitism whatever the age of the eggs or the larvae present on the host. The least avoidance was observed when the hosts were parasitized by E. vuilleti 30 min beforehand. This avoidance behavior is adaptive and is related to the low survival chances of the D. basalis larvae when they are in interspecific competition with E. vuilleti larvae. The analysis of the behavior of D. basalis demonstrated that the avoidance of multiparasitism could be due to the perception of two signals; an external signal deposited on the surface of the seeds during the E. vuilleti oviposition phase and an internal signal due to the presence of the eggs and larvae at the surface of the hosts. E. vuilleti females did not avoid multiparasitism and multiparasitized the hosts bearing D. basalis eggs or larvae. The behavior of E. vuilleti females was not disturbed by the presence of its competitor. Under these conditions of interspecific competition, the survival chances of E. vuilleti larvae were very high whatever the age of its competitor D. basalis. The two species of parasitoids could move in a column containing healthy seeds of V. unguiculata and patches with seeds containing parasitized or unparasitized larvae. The distribution of D. basalis females introduced into these columns depended on the host quality. They avoided the patches containing the hosts parasitized E. vuilleti and were found in the patches with healthy hosts. The behavior of E. vuilleti females was very different; the distribution of the females and the parasitism and multiparasitism rates were not affected by the quality of the hosts present in the patches. The adaptive significance of the behaviors of these two species was analyzed in relation to the survival chances of their offspring.     

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.     

Repeatability of Foreign Egg Rejection: Testing the Assumptions of Co‐Evolutionary Theory

Most theoretical models of coevolution between brood parasites, whether interspecific or conspecific, and their hosts explicitly assume consistent individual behaviour in host egg‐rejection responses. Accordingly, hosts cast as acceptors always accept, whereas ejectors always reject parasitic eggs when exposed to stable ecological conditions. To date, only few studies have attempted to test this critical assumption of individual repeatability in egg‐rejection responses of hosts. Here, we studied the repeatability of egg rejection in blackbirds (Turdus merula) and song thrush (T. philomelos), species in which females are reported to reject simulated, non‐mimetic foreign eggs at intermediate frequencies at the population level. However, intermediate rates of acceptance and rejection can be consistent with either or both intra‐ and interindividual variability in rejection behaviours. Our experiments revealed generally high individual consistency in these hosts’ responses to experimentally introduced non‐mimetic and mimetic model foreign eggs. Individuals also responded faster on average to second than to first trials within the same breeding attempts, but the difference was statistically significant only in blackbirds. These results are consistent with the critical assumption of co‐evolutionary models, that statistically egg rejection is mostly individually repeatable, but also reveal that some individuals in both species change their responses even within the short time‐window of one breeding attempt. The data suggest that individuals reject foreign eggs faster when perceived parasitism risk is greater because of repeated introductions of experimental parasitic eggs. We provide methodological recommendations to facilitate experimental and meta‐analytical studies of individual egg rejection repeatability and discuss how to reduce technical constraints arising from disparate treatments and variable sample sizes for future studies.     

Within-plant vertical distributions of the scale insect Nipponaclerda biwakoensis and its five parasitoids that exhibit frequent successful multiparasitism on the common reed

Within‐shoot vertical distributions of the scale insect Nipponaclerda biwakoensis, five species of gregarious parasitoid wasps attacking the scale, and successful multiparasitism (emergence of multiple parasitoid species from a single host) by the parasitoids on the common reed were investigated. Each reed shoot collected was longer than 2.0 m, and was divided into pieces of 0.5 m each from the base of the shoot. The mean number of adult female scales per 0.5 m of shoot increased with height, but the mean rate of overall parasitism of the female scales decreased with height. The five parasitoids showed species‐specific within‐shoot distribution patterns, with respect to the rate of parasitism: Astymachus japonicus, Boucekiella depressa and Encyrtidae sp. 2 showed the greatest rates of parasitism at heights of 0–0.5, 0.5–1.0 and 1.0–1.5 m, respectively. The parasitism rate by Aprostocetus sp. was higher at the lower half of the shoots, whereas that by Encyrtidae sp. 1 was higher at the central positions. The distribution of the five parasitoids largely overlapped with each other at the level of scale aggregations. Nevertheless, successful multiparasitism occurred frequently only in several types of species combinations that involved two parasitoid species showing similar within‐shoot distribution patterns. The rate of successful multiparasitism by B. depressa with A. japonicus or Aprostocetus sp., and that by Encyrtidae sp. 2 with Encyrtidae sp. 1 was constantly high at different vertical positions on the shoots. This may be because B. depressa and Encyrtidae sp. 2 preferentially oviposit into scales previously parasitized by these other species.     

The importance of nest‐site and habitat in egg recognition ability of potential hosts of the Common Cuckoo Cuculus canorus

The intensity of selection exerted by brood parasites on their hosts depends on the proportion of nests that are parasitized and the fitness costs of parasitism. Nest detection by brood parasites influences the probability of parasitism, and we propose that the difficulty faced by brood parasites of finding nests on the ground may make ground‐nesting species subject to lower levels of parasitism, causing a reduction in levels of defence compared with species breeding in shrubs, trees and elsewhere above the ground. We tested the prediction that the rejection rate of Common Cuckoo Cuculus canorus eggs by hosts is inversely related to the frequency with which they build nests on the ground, both at local and at continental scales. First, we used estimates of the rejection rate of non‐mimetic model eggs experimentally introduced into the nests of 26 potential host species breeding in the Sierra Nevada Mountains of southern Spain. Most species tested in the Sierra Nevada showed high rejection rates of both mimetic and non‐mimetic eggs, whereas the European Robin Erithacus rubecula, with a low rejection rate, was the only species that was regularly parasitized. At the continental scale we used all available published information on rejection rates of non‐mimetic models by European hosts of the Common Cuckoo. The frequency of ground‐nesting explained interspecific variation in rejection rate of non‐mimetic model eggs both for the species tested in the Sierra Nevada and for all European hosts after controlling for all other life‐history variables known to affect rejection rates. An effect of the abundance of trees in a particular habitat, previously shown to affect parasitism by the Common Cuckoo, was only apparent from analyses of continental‐scale data and not from the Sierra Nevada mountains, suggesting that particular properties of mountainous areas affect Common Cuckoo parasitism. Ground‐nesting species showed lower rejection rates than species breeding in bushes or trees. These results suggest that species nesting on the ground may have suffered lower parasitism pressures in their historical coevolutionary interactions with the Common Cuckoo.     

Intrinsic competition and its effects on the survival and development of three species of endoparasitoid wasps

In natural systems, pre‐adult stages of some insect herbivores are known to be attacked by several species of parasitoids. Under certain conditions, hosts may be simultaneously parasitized by more than one parasitoid species (= multiparasitism), even though only one parasitoid species can successfully develop in an individual host. Here, we compared development, survival, and intrinsic competitive interactions among three species of solitary larval endoparasitoids, Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae), Microplitis demolitor Wilkinson, and Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), in singly parasitized and multiparasitized hosts. The three species differed in certain traits, such as in host usage strategies and adult body size. Campoletis sonorensis and M. demolitor survived equally well to eclosion in two host species that differed profoundly in size, Pseudoplusia includens (Walker) and the larger Heliothis virescens (Fabricius) (both Lepidoptera: Noctuidae). Egg‐to‐adult development time in C. sonorensis and M. demolitor also differed in the two hosts. Moreover, adult body mass in C. sonorensis (and not M. demolitor) was greater when developing in H. virescens larvae. We then monitored the outcome of competitive interactions in host larvae that were parasitized by one parasitoid species and subsequently multiparasitized by another species at various time intervals (0, 6, 24, and 48 h) after the initial parasitism. These experiments revealed that M. croceipes was generally a superior competitor to the other two species, whereas M. demolitor was the poorest competitor, with C. sonorensis being intermediate in this capacity. However, competition sometimes incurred fitness costs in M. croceipes and C. sonorensis, with longer development time and/or smaller adult mass observed in surviving wasps emerging from multiparasitized hosts. Our results suggest that rapid growth and large size relative to competitors of a similar age may be beneficial in aggressive intrinsic competition.     

Overwintering strategy of endoparasitoids in Chilo suppressalis: a perspective from the cold hardiness of a host

Although parasitoids ultimately kill their host, koinobiont parasitoids must protect not only themselves but also their hosts against extreme environments. In this study, the parasitism rate of Chilo suppressalis Walker (Lepidoptera: Pyralidae) was investigated, and the average body weights, supercooling points, and concentrations of glycerol (acting as a cryoprotectant) in the hemolymph were compared between parasitized and non‐parasitized larvae. Five species of koinobiont endoparasitoids parasitized the overwintering C. suppressalis larvae and the total parasitism rate was 47.6% (n = 1 537). Average body weight of parasitized larvae was significantly lower than that of non‐parasitized larvae, and the parasitism rate of the lighter group (20–30 mg) was highest. The supercooling point of parasitized C. suppressalis larvae (?15.7 ± 0.3 °C) was significantly lower than that of the non‐parasitized larvae (?14.3 ± 0.2 °C). In addition, supercooling points were not correlated with body weights between parasitized and non‐parasitized larvae, indicating that cold hardiness of parasitized larvae was enhanced by endoparasitoids. Furthermore, the concentration of glycerol in the hemolymph was significantly higher in parasitized larvae (205.0 ± 7.1 μmol ml^?1) than in non‐parasitized larvae (169.8 ± 14.4 μmol ml^?1), which suggests that the mechanism that decreases the supercooling point of parasitized larvae was associated with glycerol. All these results indicated that the cold hardiness of parasitized C. suppressalis larvae was enhanced by their endoparasitoids, which benefitted overwintering endoparasitoids.     

Parasitism and venom of ectoparasitoid Scleroderma guani impairs host cellular immunity

Venom is a prominently maternal virulent factor utilized by parasitoids to overcome hosts immune defense. With respect to roles of this toxic mixture involved in manipulating hosts immunity, great interest has been mostly restricted to Ichneumonoidea parasitoids associated with polydnavirus (PDV), of which venom is usually considered as a helper component to enhance the role of PDV, and limited Chalcidoidea species. In contrast, little information is available in other parasitoids, especially ectoparasitic species not carrying PDV. The ectoparasitoid Scleroderma guani injects venom into its host, Tenebrio molitor, implying its venom was involved in suppression of hosts immune response for successful parasitism. Thus, we investigated the effects of parasitism and venom of this parasitoid on counteracting the cellular immunity of its host by examining changes of hemocyte counts, and hemocyte spreading and encapsulation ability. Total hemocyte counts were elevated in parasitized and venom‐injected pupae. The spreading behavior of both granulocytes and plasmatocytes was impaired by parasitization and venom. High concentration of venom led to more severely increased hemocyte counts and suppression of hemocyte spreading. The ability of hemocyte encapsulation was inhibited by venom in vitro. In addition to immediate effects observed, venom showed persistent interference in hosts cellular immunity. These results indicate that venom alone from S. guani plays a pivotal role in blocking hosts cellular immune response, serving as a regulator that guarantees the successful development of its progenies. The findings provide a foundation for further investigation of the underlying mechanisms in immune inhibitory action of S. guani venom.     

Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

Phenology of Parasetigena silvestris (Diptera: Tachinidae), gypsy moth (Lymantria dispar) (Lepidoptera: Lymantriidae) larval parasitoid and its efficiency for parasitisation

Parasetigena silvestris is a univoltine, solitary, larval endoparasitoid which lays its eggs on the surface of gypsy moth larvae. Field collection of the host larvae (2nd through 5th instar) from an artificially established gypsy moth population were made to compare stage specific parasitism between larvae without and with P. silvestris tachinid eggs. The tachinid oviposition rate detected was highest in second instar larvae, and then decreased as larvae developed toward full maturity. The opposite was true for tachinid parasitoid emergence which had no emergence from second through third host instar larvae. Fourth instar gypsy moth larvae, however, experienced significantly higher parasitism by P. silvestris in the larvae with eggs than those without the eggs. The braconid wasp Cotesia melanoscelus caused significantly higher parasitism in early instar larvae with P. silvestris eggs than in those without the eggs. The tachinid prefers to lay more eggs on parasitised larvae by the braconid even though the braconid is a superior competitor to the fly during multiparasitism. Factors influencing parasitism rates by P. silvestris such as host-parasitoid synchronisation and the multiparasitism interaction with C. melanoscelus are discussed.     

Reproductive strategies under multiparasitism in natural populations of the parasitoid wasp Nasonia (Hymenoptera)

Parasitoid Nasonia wasps adjust their progeny sex ratio to the presence of conspecifics to optimize their fitness. Another trait under female control is the induction of offspring diapause. We analysed progeny sex ratios and the proportion of diapausing offspring of individual Nasonia females in host patches parasitized by two species, Nasonia vitripennis and Nasonia giraulti, in North American field populations using microsatellite fingerprinting. Both Nasonia species produced similar sex ratios on hosts that were co‐parasitized by their own species as by the other species, indicating that females do not distinguish between con‐ and heterospecific clutches. The sex ratios of the diapause and adult fractions of mixed broods from single females were not correlated. We found further indications that N. vitripennis females take the emergence time of the offspring into account in their sex allocation. The reproductive strategies of Nasonia under multiparasitism are largely adaptive, but also partially constrained by information.     

Intra- and interspecific competition by Fopius arisanus and Diachasmimorpha tryoni (Hymenoptera: Braconidae), parasitoids of tephritid fruit flies

Fopius arisanus (Sonan) and Diachasmimorpha tryoni (Cameron) are two important solitary endoparasitoids of tephritid fruit flies. The former species attacks host eggs while the latter attacks host larvae, and both species emerge as adults from the host puparium. This study investigated intrinsic competition between these two parasitoids, as well as aspects of intraspecific competition within each species in the Mediterranean fruit fly, Ceratitis capitata (Wiedemann). Parasitization by F. arisanus resulted in direct mortality of host eggs and prolonged development of host eggs and larvae. Superparasitism by F. arisanus was uncommon when mean parasitism per host patch was <50%, but increased with rising rates of parasitism. Superparasitism by D. tryoni was more common. In superparasitized hosts, supernumerary individuals of F. arisanus were killed through physiological suppression, while supernumerary larvae of D. tryoni were killed mainly through physical attack. In multiparasitized hosts, dissections showed that 81.6% of D. tryoni eggs in the presence of F. arisanus larvae died within 3 days, indicating physiological inhibition of egg hatch. Rearing results further showed that F. arisanus won almost all competitions against D. tryoni. The ratio of D. tryoni stings to ovipositions was lower in hosts not previously parasitized by F. arisanus than in parasitized hosts, suggesting that D. tryoni can discriminate against parasitized hosts. The mechanism that F. arisanus employs to eliminate D. tryoni is similar to that it uses against all other larval fruit fly parasitoids so far reported. The results are discussed in relation to the competitive superiority of early acting species in fruit fly parasitoids, and to a possible competitive-mediated mechanism underlying host shift by D. tryoni to attack non-target flies following the successful introduction of F. arisanus in Hawaii.     

Tritrophic interactions and trade-offs in herbivore fecundity on hybridising host plants

Abstract. 1. Interspecific plant hybridisation can have important evolutionary consequences for hybridising plants and for the organisms that they interact with on multiple trophic levels. In this study the effects of plant hybridisation on the abundance of herbivores and on the levels of herbivore parasitism were investigated. 2. Borrichia frutescens, B. arborescens, and their hybrid (B. × cubana) were censused for Asphondylia borrichiae galls and Pissonotus quadripustulatus plant hoppers in the Florida Keys. Levels of egg parasitism were determined by dissecting parental and hybrid stems and galls for herbivore and parasite eggs and larvae. Stem toughness and gall size are plant‐mediated modes of protection from parasitism and these were also measured. For gall midges, fly size was measured as an estimate of fecundity. 3. Field censuses indicated that herbivore abundances varied on hybrid hosts relative to parent plant species and that the different herbivore species exhibited different patterns of abundance. Asphondylia borrichiae gall numbers followed the additive pattern of abundance while P. quadripustulatus numbers most closely resembled the dominance pattern. 4. Parasitism of P. quadripustulatus eggs was high on B. frutescens and the hybrids, and low on B. arborescens, which also had significantly tougher stems. Asphondylia borrichiae suffered the highest levels of parasitism on B. frutescens, the host plant which produced the smallest galls. On B. arborescens, which produced the largest galls, levels of A. borrichiae parasitism were lowest. Both parasitism and gall size were intermediate on the hybrid plants. Galls on B. arborescens and hybrid plants produced significantly smaller flies then those from B. frutescens suggesting that, when selecting hosts from among parent species and hybrids, gall flies may face a trade‐off between escape from natural enemies and maximising fecundity.     

Egg Rejection and Brain Size among Potential Hosts of the Common Cuckoo

Interspecific brood parasitism by the common cuckoo (Cuculus canorus) lowers host fitness, and has selected for discrimination and rejection of parasitic eggs in their commonly parasitized hosts. Cognitive demands needed to discriminate and reject cuckoo eggs may have led to augmentation of relative brain size among passerine hosts parasitized by cuckoos. This hypothesis predicts for across species positive relationships of brain size with rejection rate, host suitability and parasitism level. Here we test these predictions while controlling for phylogenetic, ecological and developmental factors known to affect brain size and egg rejection in a comparative study using the cuckoo and their hosts in Europe as a model system. Contrary to expected the rate of rejection of non‐mimetic cuckoo eggs covaried negatively with relative brain size across bird species. Either suitability as cuckoo host, which reflects long‐time duration of exposure to cuckoo parasitism, and level of parasitism, did not relate to brain size. Our results do not support the hypothesis that cuckoo parasitism was a main direct force affecting brain size variation across passerine hosts.