Host-Parasitoid Dynamics and the Success of Biological Control When Parasitoids Are Prone to Allee Effects

In sexual organisms, low population density can result in mating failures and subsequently yields a low population growth rate and high chance of extinction. For species that are in tight interaction, as in host-parasitoid systems, population dynamics are primarily constrained by demographic interdependences, so that mating failures may have much more intricate consequences. Our main objective is to study the demographic consequences of parasitoid mating failures at low density and its consequences on the success of biological control. For this, we developed a deterministic host-parasitoid model with a mate-finding Allee effect, allowing to tackle interactions between the Allee effect and key determinants of host-parasitoid demography such as the distribution of parasitoid attacks and host competition. Our study shows that parasitoid mating failures at low density result in an extinction threshold and increase the domain of parasitoid deterministic extinction. When proned to mate finding difficulties, parasitoids with cyclic dynamics or low searching efficiency go extinct; parasitoids with high searching efficiency may either persist or go extinct, depending on host intraspecific competition. We show that parasitoids suitable as biocontrol agents for their ability to reduce host populations are particularly likely to suffer from mate-finding Allee effects. This study highlights novel perspectives for understanding of the dynamics observed in natural host-parasitoid systems and improving the success of parasitoid introductions.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Demonstration of interspecific competition between two sympatric egg parasitoids of Riptortus pedestris (Fabricius) in laboratory condition

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) acts as a facultative hyperparasitoid of Gryon japonicum (Ashmead) (Hymenoptera: Platygastridae) sympatric parasitoid of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae). A longer period of host egg exploitation by both parasitoid species would be beneficial for O. nezarae, while G. japonicum tends to be successful when the parasitoids have only a short exploitation period. We demonstrated the interspecific competition by measuring parasitism in nine combinations of host densities (10, 20, and 40 eggs) and exploitation times (1, 3, and 5 days). To reflect the gregarious-solitary dichotomy of the two species, three O. nezarae and one G. japonicum mated females were compared in addition to a one-to-one competition design. We found that O. nezarae was the better competitor when exploitation time was longer than 1 day, irrespective of host density. Total parasitism rate and progeny emergence of O. nezarae were 1.6–2.8 and 4.7–7.3 times higher than for G. japonicum in three-to-one competition design, respectively. Although G. japonicum females were more effective in host finding (as shown by their higher per capita rate of parasitism when exploitation time was short), their progeny suffered high mortality from the larval interspecific competition inside multiparasitized host eggs. These results suggest that gregarious O. nezarae is the superior competitor when host eggs are available for longer period of time while solitary G. japonicum is superior when host resources are available for only a limited time.     

Does parasitoid state affect host range expression?

The pre-release risk assessment of parasitoids for classical biological control generally involves non-target testing to define the agent’s host range. To ensure that no suitable host species are falsely rejected in these tests, it has been suggested that the physiological and informational state of parasitoids be manipulated to enhance their “motivation to oviposit”. However, the effects of such factors on host acceptance are not consistent across parasitoid species, making it laborious to identify the conditions necessary to maximise host acceptance. Our objective was to determine whether changes in parasitoid state could alter host acceptance behaviour sufficiently to affect host range expression. In addition, we tested the assumption that a state-dependent shift in motivation to oviposit on the target host will translate to a similar change in responsiveness to lower-ranked host species. Three-day-old and 10-day-old females of the candidate classical biological control agent, Diadromus pulchellus, were offered 12 non-target species of varying relatedness to the target pest, Acrolepiopsis assectella, in a series of no-choice and choice oviposition trials. Younger D. pulchellus females had previously demonstrated greater motivation to oviposit in the target pest and were, therefore, predicted to express a broader host range than older females. Parasitoid age had a minor effect on host range expression that was contrary to expectations. Older females more readily attacked one of the non-target species in no-choice tests and inflicted higher mortality in one of the choice tests. Ultimately however, young and old parasitoids still attacked the same four non-target species and their offspring emerged from the same three. There was an interaction between the effects of parasitoid condition and experimental design on responsiveness to low-ranked hosts: increasing non-target density in choice tests significantly altered attack rates by 10-day-old, but not by 3-day-old, parasitoids. The implications of these findings for host specificity testing depend largely on the specific aims of a host range assessment. Parasitoid state influenced the frequency of non-target attack but did not affect which non-target species were attacked.     

Effects of floral resources on the efficacy of a primary parasitoid and a facultative hyperparasitoid

Resources added to agroecosystems to enhance biological control are potentially available to multiple members of the resident insect community—not only the biological control agents for which the resources are intended. Many studies have examined the effects of sugar feeding on the efficacy of biological control agents. However, such information is lacking for other, interacting species such as facultative hyperparasitoids, which may contribute to pest suppression but can also interfere with introduced biological control agents. Under greenhouse conditions, we tested the direct effects of sugar and nectar provisioning on the longevity, host‐killing impact and offspring production of two pupal parasitoids associated with leek moth, Acrolepiopsis assectella: the introduced biological control agent, Diadromus pulchellus, and the native facultative hyperparasitoid, Conura albifrons. Adding sucrose, buckwheat or a combination of buckwheat and common vetch to a sugar‐deprived system (potted leek plants in cages) increased parasitoid longevity and resulted in higher leek moth parasitism and mortality compared to water or common vetch treatments. However, the two parasitoid species exhibited a distinct temporal response to the treatments, likely influenced by differences in their life histories. This study provides insight into how integrating conservation biological control techniques could affect the success of a classical biological control programme.     

The role of volatiles from cruciferous plants and pre-flight experience in the foraging behaviour of the specialist parasitoid Cotesia plutellae

The braconid Cotesia plutellae is an important larval parasitoid of the diamondback moth (Plutella xylostella), a major pest of crucifers in the tropics and sub-tropics. The in-flight searching behaviour of C. plutellae was investigated in a wind tunnel and the close-range attack behaviour observed in cages. The relative importance of volatile stimuli emanating from the plant-host-complex, oilseed rape (Brassica napus) – P. xylostella, in the long-range attraction of C. plutellae was investigated. Plants that were mechanically damaged, or damaged by P. xylostella larvae, were attractive to the parasitoid. Host-damaged leaves remained attractive to the parasitoid after removal of the host larvae. These results indicate that C. plutellae predominantly uses plant derived stimuli in its in-flight searching behaviour. An oviposition experience or contact with a host-damaged leaf prior to the bioassay significantly increased the response to these volatile cues. The foraging behaviour of C. plutellae is compared with other braconid larval parasitoids attacking lepidopteran hosts on crucifers.     

Potential impact of the native hyperparasitoid Conura albifrons (Hymenoptera: Chalcididae) on the exotic biological control agent Diadromus pulchellus (Hymenoptera: Ichneumonidae)

The solitary pupal parasitoid Diadromus pulchellus was released in 2010 as a classical biological control agent against leek moth, Acrolepiopsis assectella, an important new pest of onion crops, Allium spp. in Eastern North America. Post-release monitoring using sentinel leek moth pupae revealed that the facultative hyperparasitoid Conura albifrons was attacking leek moth and potentially D. pulchellus as well. We used laboratory choice and no-choice tests to assess the potential impact of C. albifrons on leek moth and D. pulchellus. C. albifrons is capable of developing in newly formed leek moth pupae and leek moth pupae containing first instar D. pulchellus, as well as in fully developed D. pulchellus pupae. Survivorship of both leek moth and D. pulchellus exposed to C. albifrons was significantly lower than that of unexposed controls. In choice trials, prior host experience significantly influenced host choice by C. albifrons. These results suggest that C. albifrons could impact the establishment of D. pulchellus through both competition and intraguild predation, and that the impact has the potential to change as the relative frequency of the two hosts shifts in field populations.     

The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Host size of Pteromalus puparum, a gregarious pupal parasitoid, shows a wide inter- and intraspecific variation. Experiments were made to study the regulation of the number and sex ratio of progeny per host by the parasitoid. The parasitoid could discriminate inter- and intraspecific size differences of the host and regulate the number of eggs according to the host size when a single female attacked the host. The sex ratio of progeny (proportion males) was about 0.1. The number of progeny laid by the female agreed with the energetically most efficient number og eggs in order to maximize total weight of progeny per host but not with the reproductively most efficient number of eggs to maximize the total fecundity of the progeny. The parasitoid laid smaller number of eggs in a half buried host, but the number was much larger than a half of those in a fully exposed host. When more than one female attacked a single host, the number and sex ratio of progeny per host increased with the number of females attacking the host, but the number of progeny per female decreased. The change of the sex ratio agreed with the prediction of the local mate competition model.     

The use of kairomones for foraging decisions by an aphid parasitoid in small host aggregations

1. The wasp Diaeretiella rapae uses honeydew emitted by its host, the cabbage aphid Brevicoryne brassica, as a kairomone (chemicals emitted by an organism as part of its activity and used by its natural enemies to their advantage). The role of the kairomone in foraging decisions by the parasitic wasp was explored by manipulating the amount of honeydew and the number of aphids in a colony independently. The count-down patch-exploitation mechanism (Iwasa et al., 1981) was employed to predict the results of these manipulations and contrast them with the predictions of Waage's (1979) model. 2. Kairomonal activity of honeydew decreased as the honeydew aged and lost its activity completely within 72 h. 3. The wasp was exposed to different amounts of honeydew on (a) aphid-free leaves and (b) leaves bearing colonies of 150 aphids. The parasitoid search time on both leaf types increased with increasing honeydew contamination. On aphid-bearing leaves, the number of attacked aphids in the colonies also increased with increasing honeydew contamination. The presence of aphids reduced the parasitoid search time compared to search time on aphid-bearing leaves with the same level of honeydew contamination, as predicted by the count-down model. 4. Parasitoids exposed to the same amount of honeydew, but a different number of aphids in a colony on the leaf, first increased the number of aphids attacked in relation to the number of aphids in the colony. But as the colony reached a certain size, the number of aphids attacked levelled off, despite the increase in the number of aphids in the colony. Search time was variable but did not exhibit any trend as the number of aphids in the colonies increased. 5. These results suggest that honeydew level is used by D. rapae as a cue for assessment of the number of aphids in the colony. In such cases, a count-down exploitation mechanism gives the best results to a forager. Many parasitoids may use kairomones for patch assessment and will therefore employ a count-down rule during patch exploitation.     

Early experience influences several steps of the host selection process differentially in Aphidius ervi (Hymenoptera,Braconidae)

Experience of host-associated olfactory stimuli during development affects subsequent foraging decisions in many parasitoids, leading to host fidelity. We have recently shown that odours emitted by an alien host-plant complex (HPC) may affect this learning process. However, the consequences of this olfactory experience on parasitoid host foraging decisions is unknown. Here, we investigated if olfactory preferences induced by experience might drive parasitoid HPC choice and oviposition decisions. We presented two HPCs in dual choice experiments to Aphidius ervi females that had been reared on each HPC, either in a simple (one HPC) or complex (two HPCs) olfactory environment. HPC choice, time before landing on a HPC and number of aphid attacks were recorded. Early experience had contrasting effects on each of these parameters. Regardless of their origin, parasitoids did not land preferentially on any HPC, but they more frequently attacked one of the two hosts (Acyrthosiphon pisum) once they had landed on its HPC. Females emerging from the A. pisum/faba bean HPC attacked more aphids, regardless of the host species. Finally, the olfactory complexity of the environment had inconsistent effects on foraging decisions. These observations, contrasting with previous results obtained in olfactometer assays, indicate that olfactory preferences induced in early stages are not sufficient to predict parasitoid oviposition decisions. Moreover, the integration of multiple signals at several scales might generate different foraging patterns at each step of the host selection process.     

Role of Caterpillar Silk Thread in Location of Host Pupae by the Parasitoid <Emphasis Type="Italic">Diadromus pulchellus</Emphasis>

Diadromus pulchellus is a solitary ichneumonid parasitoid. Its only known host is the pupa of Acrolepiopsis assectella, a specialist herbivore of Allium species. D. pulchellus females parasitize A. assectella pupae within 48 h after the caterpillars spin their cocoon and begin to pupate. Having observed that the cocoon produced by the leek moth caterpillar stimulates parasitoid egg-laying and that caterpillar leaves a silk thread, we studied the hypothesis that silk thread might be involved in host-finding by the parasitoid. Behavioral tests showed that when D. pulchellus females encounter a host silk thread, they change directions, follow the thread, and quickly locate the host. These findings show that pupal parasitoids can use signals produced by their hosts at the developmental instar preceding the one that they parasitize.     

Competitive Interactions between Parasitoids Provide New Insight into Host Suppression

Understanding the dynamics of potential inter- and intraspecific competition in parasitoid communities is crucial in the screening of efficient parasitoid species and for utilization of the best parasitoid species combinations. In this respect, the host-parasitoid systems, Bemisia tabaci and two parasitoids, Eretmocerus hayati (exotic) and Encarsia sophia (existing) were studied under laboratory conditions to investigate whether interference competition between the exotic and existing species occurs as well as the influence of potential interference competition on the suppression of the host B. tabaci. Studies on interspecific-, intraspecific- and self-interference competition in two parasitoid species were conducted under both rich and limited host resource conditions. Results showed that (1) both parasitoid species negatively affect the progeny production of the other under both rich and limited host resource conditions; (2) both parasitoid species interfered intraspecifically on conspecific parasitized hosts when the available hosts are scarce and; 3) the mortality of B. tabaci induced by parasitoids via parasitism, host-feeding or both parasitism and host-feeding together varied among treatments under different host resource conditions, but showed promise for optimizing control strategies. As a result of our current findings, we suggest a need to investigate the interactions between the two parasitoids on continuous generations.     

Combined effects of host quality and local mate competition on sex allocation inLariophagus distinguendus

Summary Many parasitoid wasps are known to adjust sex ratio in response to either local mate competition (LMC) or host quality. Nevertheless, few studies have investigated the combined effects of these two factors on sex allocation. The sex allocation pattern inLariophagus distinguendus, a parasitoid of granary weevil larvae, is contrasted to the expectations of Werren's (1984) model combining LMC and host quality. Several predictions of the model are confirmed, but others are not. Sex ratio on both large and small hosts declines with proportion of small hosts attacked in a manner consistent with the model. However, when only one host size is parasitized, sex ratio is not independent of that host size, as predicted by the model. Various possibilities for the deviation between expected and observed are discussed. A partial LMC/host quality model is developed which allows for some matings outside the natal patch, and predictions of this model conform more closely to the pattern observed inL. distinguendus. Finally, the application of parasitoid studies to basic questions in evolutionary ecology is briefly discussed.     

The effects of patch richness on con-specific interference in the parasitoid Ibalia leucospoides (Hymenoptera: Ibaliidae)

Abstract We experimentally studied con-specific interference in Ibalia leucospoides, a parasitoid of the woodwasp Sirex noctilio, on pine logs containing variable numbers of woodwasp larvae. Competiton occurred when two different-sized female parasitoids foraged on a host patch, and consequently the small female always abandoned the patch. Regardless of host density, con-specific presence did not affect the attack rate, the number of hosts attacked, nor patch residence time by the winner, when compared to a control female that foraged alone. In contrast, con-specific presence reduced patch time and the number of hosts attacked by the loser. Finally patch time (by both) as well as number of hosts attacked (by the winner only) increased with host density per patch. Our results suggest that con-specific presence has different consequences for different-sized females during patch exploitation.     

Beneficial arthropod behavior mediated by airborne semiochemicals. VIII. Learning of host-related odors induced by a brief contact experience with host by-products inCotesia marginiventris (Cresson), a generalist larval parasitoid

Females of Cotesia marginiventris(Cresson), a generalist larval parasitoid, were observed to respond to host related odors in a four-arm olfactometer. The females were significantly more responsive to the odors after a brief contact experience with host-damaged leaves contaminated with host by products. During the experience, actual encounters with hosts were not required to improve subsequent responses to host-related odors. The response to odors of the plant-host complex with which parasitoids had experience was significantly higher than the response to odors of an alternative plant-host complex. This suggests that the experience effect is due, at least partly, to associative learning. We suspect that females of this generalist parasitoid, as was recently found for those of a specialist, recognize specific semiochemicals when they contact frass of suitable host larvae. The parasitoids, subsequently, associate the surrounding odors with the possible presence of hosts, and use these odors as cues in their search for more hosts. This could be an important component in the host-searching behavior of many parasitoids.Mention of a proprietary product does not constitute an endorsement or the recommendation for its use by the USDA.     

Temporal/spatial structure and the dynamical property of laboratory host-parasitoid systems

The effects of spatial structure in terms of local capacity, or the maximum number of larvae surviving competition at resource patches, and temporal structure in terms of the period vulnerable to parasitoid attack in host populations on the persistence of host-parasitoid systems were quantitatively evaluated by laboratory experiments and well-parameterized model analyses. One of two bruchid beetles,Callosobruchus maculatus andC. phaseoli, were used as a host with Heterospilus prosopidis used as the parasitoid.C. maculatus, in which few larvae survive competition to become adults in each bean, andC. phaseoli, in which many larvae become adults in each bean, along with two kinds of beans, the mung and the azuki, were combined to construct four (2×2) resource-herbivorous host-parasitoid systems that differed in local capacity and vulnerable period. The mung-C. maculatus system with the parasitoid was the most persistent, i.e., took the longest time for extinction of either the host or parasitoid to occur. Since this resource-herbivorous host combination exhibited the lowest local capacity and the shortest vulnerable period, these two conditions possibly promoted the persistence of the system. A model incorporating the host population structure supported the observed persistence. Furthermore, the possible contribution of the timing of density-dependent competition of the host on the host-parasitoid persistence is predicted.     

Pea aphid clonal resistance to the endophagous parasitoid Aphidius ervi

The physiological mechanism of resistance to the endophagous braconid Aphidius ervi Haliday (Hymenoptera, Braconidae) by a pink clone (PC) of Acyrthosiphon pisum (Harris) (Homoptera, Aphididae) has been investigated. Comparative data on parasitoid development and associated host biochemical changes in the resistant PC aphids and in a susceptible green clone (GC) of A. pisum are reported. When the PC aphids were attacked as early 4th instars, the developing parasitoid larvae showed a strongly reduced increase in size, compared to those synchronously developing in GC aphids, and were unable to produce a regular mummy. In contrast, parasitism of 2nd instar PC aphids, allowed completion of parasitoid development, but adults had a prolonged developmental time, due to a longer duration of parasitoid’s final (3rd) instar. In all cases, teratocytes, cells deriving from the A. ervi serosal membrane, and the proteins abundantly synthesised by them, were never found in the haemolymph of parasitised PC aphids. Host castration, as demonstrated by total protein incorporation into reproductive tissues, was total in the majority of early (2nd instar) parasitised host aphids, while it was limited when later instars (4th) of PC aphids were parasitised. This is partly due to the absence of the cytolytic activity of teratocytes on host embryos, which, through their persistence, may compete for nutritional resources with the developing parasitoid larvae. In parasitised PC aphids, this competitive effect is further aggravated for the parasitoid by the absence of the regulated amino acid titre increase in the host haemolymph, which is regularly observed in GC aphids. Failure of teratocyte development in the PC clone of the pea aphid is, then, the major functional constraint accounting for the reduction/inhibition of A. ervi larval growth. The reported results allow to assess in vivo the role of teratocytes in the host physiological redirection and nutritional exploitation by the parasitoid, and to integrate and validate the proposed physiological model of host-parasitoid interactions in the system A. pisum-A.ervi.     

Fitness costs of intrinsic competition in two egg parasitoids of a true bug

Intrinsic competition in insect parasitoids occurs when supernumerary larvae develop in the same host as consequence of multiple ovipositions by females of the same species (intra-specific competition) or by females of different species (inter-specific competition). Studies on intrinsic competition have mainly focused on understanding the factors that play a role in the outcome of competition, while fitness-related effects for the parasitoid surviving the competition have been poorly investigated, especially in egg parasitoids. Interestingly, even the winning parasitoid can experience fitness costs due to larval development in a host in which multiple factors have been injected by the ovipositing females or released by their larvae. In this paper we studied fitness-related traits associated with intra- and inter-specific competition between Trissolcus basalis (Wollaston) and Ooencyrtus telenomicida (Vassiliev), the main egg parasitoids associated with the southern green stink bug Nezara viridula (L.) in Italy. We investigated the impact of intrinsic competition for the surviving parasitoid in terms of body size, developmental time, number and size of oocytes. Our results indicated that T. basalis adults did not experience fitness-related costs when surviving intra-specific competition; however, adults were smaller, took longer to develop and females produced fewer oocytes after surviving inter-specific competition. A different outcome was found for O. telenomicida where the emerging females were smaller, produced fewer and smaller oocytes when suffering intra-specific competition whereas no fitness costs were found when adults survived inter-specific competition. These results support the hypothesis that the impact of intrinsic competition in egg parasitoids depends on the severity of the competitive interaction, as fitness costs were more pronounced when the surviving parasitoid interacted with the most detrimental competitor.     

Interspecific competition between Diadegma semiclausum Hellen and Diadegma mollipla (Holmgren), parasitoids of the diamondback moth, Plutella xylostella (L), feeding on a new host plant

Interspecific competition between an introduced parasitoid species aimed at controlling a herbivorous pest species and a native parasitoid parasitising the same host may influence the success of classical biological control programmes. In Kenya, interspecific competition between an introduced and a local parasitoid on two diamondback moth populations (DBM, Plutella xylostella) was investigated on two different host plants. We tested simultaneous and delayed competition of the local parasitoid Diadegma mollipla Holmgren and its exotic congenus D. semiclausum Hellen on a newly aquired DBM host plant (snowpea) in the laboratory. Under simultaneous competition, D. mollipla produced more progeny than D. semiclausum on snowpea. A head start of D. Mollipla, of four and eight hours before its congenus was introduced, resulted in a similar number of progeny of both species. In delayed competition (time intervals of 24 h, 48 h and 72 h), progeny production was similar for both parasitoids when the time interval was 24 h, irrespective of which species parasitized first. More progeny was produced by the species which attacked first, when the time interval was greater than 24 h, although it was only significant at 72 h. Competitive abilites of both parasitoids on the new host plant differed largely between laboratory and semi-field conditions. The influence of two host plants (snowpea and cabbage) on competition was studied in the greenhouse with different host and parasitoid densities. Parasitism levels of D. semiclausum were significantly higher than those of D. mollipla, regardless of host plant, host and parasitoid densities, but progeny production of D. mollipla on snowpea was still slightly higher than on cabbage. As compared to the confinement of parasitoids and larvae to small containers, D. mollipla parasitized very few larvae in the cages. Competitive ability of the two parasitoid species tested was influenced both by the density of the searching females and by parameters related to either the host plant and/or the herbivorous hosts.     

The impact of parasitoid emergence time on host–parasitoid population dynamics

We investigate the effect of parasitoid phenology on host–parasitoid population cycles. Recent experimental research has shown that parasitized hosts can continue to interact with their unparasitized counterparts through competition. Parasitoid phenology, in particular the timing of emergence from the host, determines the duration of this competition. We construct a discrete-time host–parasitoid model in which within-generation dynamics associated with parasitoid timing is explicitly incorporated. We found that late-emerging parasitoids induce less severe, but more frequent, host outbreaks, independent of the choice of competition model. The competition experienced by the parasitized host reduces the parasitoids’ numerical response to changes in host numbers, preventing the ‘boom-bust’ dynamics associated with more efficient parasitoids. We tested our findings against experimental data for the forest tent caterpillar (Malacosoma disstria Hübner) system, where a large number of consecutive years at a high host density is synonymous with severe forest damage.