When will rejection of parasite nestlings by hosts of nonevicting avian brood parasites be favored? A misimprinting-equilibrium model

It has been suggested that discrimination and rejection of thenestlings of avian brood parasites are most likely to evolvewhen the parasite nestling is raised alongside the host nestlings,for example, many cowbird-host systems. Under these circumstances,the benefits of discrimination are high because the host parentsmay save most of their brood. However, there is a general absenceof nestling rejection behavior among hosts of nonevicting parasites.In a cost-benefit equilibrium model, based on the premise thathost species learn to recognize their offspring through imprintingon first breeding, we show that nestling recognition can beadaptive for hosts of cowbirds, but only under strict conditions.Namely, when host nestling survival alongside the parasite islow, rates of parasitism are high and the average clutch sizeis large. All of these conditions are seldom simultaneouslyachieved in real systems. Most importantly, the parasite nestling,on average, does not sufficiently depress host nestling survivalto outweigh the costs of nestling recognition and rejectionerrors. Thus, we argue that nestling acceptance behaviors byhosts of nonevicting brood parasites may be explained as anevolutionary equilibrium in which recognition costs act as astabilizing selection pressure against rejection when most ofthe host's offspring survive parasitism.     

Coevolution of Contrary Choices in Host-Parasitoid Systems

We investigate patch selection strategies of hosts and parasitoids in heterogeneous environments. Previous theoretical work showed that when host traits vary among patches, coevolved populations of hosts and parasitoids make congruent choices (i.e., hosts and parasitoids preferentially select the same patches) and exhibit direct density dependence in the distribution of percent parasitism. However, host-parasitoid systems in the field show a range of patterns in percent parasitism, while behavioral studies indicate that hosts and parasitoids can exhibit contrary choices (i.e., hosts avoid patches favored by the parasitoid). We extend previous theory by permitting life-history traits of the parasitoid as well as the host to vary among patches. Our analysis implies that in coevolutionarily stable populations, hosts preferentially select patches that intrinsically support higher host equilibrium numbers (i.e., the equilibrium number achieved by hosts when both populations are confined to a single patch) and that parasitoids preferentially select patches that intrinsically support higher parasitoid equilibrium numbers (i.e., the equilibrium number achieved by the parasitoids when both populations are confined to a patch). Using this result, we show how variation in life-history traits among patches leads to contrary or congruent choices or leads to direct density dependence, inverse density dependence, or density independence in the distribution of percent parasitism. In addition, we determine when populations playing the coevolutionarily stable strategies are ecologically stable. Our analysis shows that heterogeneous environments containing patches where the intrinsic rate of growth of the host and the survivorship rate of the parasitoid are low result in the coevolved populations exhibiting contrary choices and, as a result, promote ecological stability.     

寄生蜂取食寄主特性及其在害虫生物防治中的作用

许多寄生性天敌昆虫的雌虫不仅寄生寄主, 而且还能取食寄主。在卵育型（synovigenic）寄生蜂类群中, 取食寄主行为是较为普遍的现象。本文综合近20年相关研究进展, 从寄生蜂类群、取食类型、生态学意义及影响因子等方面对寄生蜂的取食寄主行为进行了归纳总结。寄生蜂通过取食不仅可以杀死寄主, 直接起到控制害虫种群数量的作用, 还能通过取食策略为卵的成熟和再生提供营养来源, 对延长雌虫的寿命也有一定的帮助。对取食寄主行为的了解可为筛选优势寄生性天敌种类、评估寄生蜂在害虫生物防治中的作用提供重要信息。     

Models for integrated pest control and their biological implications

Successful integrated pest management (IPM) control programmes depend on many factors which include host-parasitoid ratios, starting densities, timings of parasitoid releases, dosages and timings of insecticide applications and levels of host-feeding and parasitism. Mathematical models can help us to clarify and predict the effects of such factors on the stability of host-parasitoid systems, which we illustrate here by extending the classical continuous and discrete host-parasitoid models to include an IPM control programme. The results indicate that one of three control methods can maintain the host level below the economic threshold (ET) in relation to different ET levels, initial densities of host and parasitoid populations and host-parasitoid ratios. The effects of host intrinsic growth rate and parasitoid searching efficiency on host mean outbreak period can be calculated numerically from the models presented. The instantaneous pest killing rate of an insecticide application is also estimated from the models. The results imply that the modelling methods described can help in the design of appropriate control strategies and assist management decision-making. The results also indicate that a high initial density of parasitoids (such as in inundative releases) and high parasitoid inter-generational survival rates will lead to more frequent host outbreaks and, therefore, greater economic damage. The biological implications of this counter intuitive result are discussed.     

Delayed action insecticides and their role in mosquito and malaria control

There is considerable interest in the management of insecticide resistance in mosquitoes. One possible approach to slowing down the evolution of resistance is to use late-life-acting (LLA) insecticides that selectively kill only the old mosquitoes that transmit malaria, thereby reducing selection pressure favoring resistance. In this paper we consider an age-structured compartmental model for malaria with two mosquito strains that differ in resistance to insecticide, using an SEI approach to model malaria in the mosquitoes and thereby incorporating the parasite developmental times for the two strains. The human population is modeled using an SEI approach. We consider both conventional insecticides that target all adult mosquitoes, and LLA insecticides that target only old mosquitoes. According to linearised theory the potency of the insecticide affects mainly the speed of evolution of resistance. Mutations that confer resistance can also affect other parameters such as mean adult life span and parasite developmental time. For both conventional and LLA insecticides the stability of the malaria-free equilibrium, with only the resistant mosquito strain present, depends mainly on these other parameters. This suggests that the main long term role of an insecticide could be to induce genetic changes that have a desirable effect on a vital parameter such as adult life span. However, when this equilibrium is unstable, numerical simulations suggest that a potent LLA insecticide can slow down the spread of malaria in humans but that the timing of its action is very important.     

Parasitism of Molluscs by Nematodes: Types of Associations and Evolutionary Trends

Although there are no confirmed fossil records of mollusc parasitic nematodes, diverse associations of more than 108 described nematode species with slugs and snails provide a fertile ground for speculation of how mollusc parasitism evolved in nematodes. Current phylogenic resolution suggests that molluscs have been independently acquired as hosts on a number of occasions. However, molluscs are significant as hosts for only two major groups of nematodes: as intermediate hosts for metastrongyloids and as definitive hosts for a number of rhabditids. Of the 61 species of nematodes known to use molluscs as intermediate hosts, 49 belong to Metastrongyloidea (Order Strongylida); of the 47 species of nematodes that use molluscs as definitive hosts, 33 belong to the Order Rhabditida. Recent phylogenetic hypotheses have been unable to resolve whether metastrongyloids are sister taxa to those rhabditids that use molluscs as definitive hosts. Although most rhabditid nematodes have been reported not to kill their mollusc hosts prior to their reproduction, some species are pathogenic. In fact, infective juveniles of Phasmarhabditis hermaphrodita vector a lethal bacterium into the slug host in which they reproduce. This life cycle is remarkably similar to the entomopathogenic nematodes in the families Steinernematidae and Heterorhabditidae. Also, the discoveries of Alloionema and Pellioditis in slugs are interesting, as these species have been speculated to represent the ancestral forms of the entomopathogenic nematodes. Development of the infective stage appears to be an important step toward the acquisition of molluscs as definitive hosts, and the association with specific bacteria may have arisen in conjunction with the evolution of necromeny.     

Mating and host density affect host feeding and parasitism in two species of whitefly parasitoids

Abstract The parasitoids in the genera of Encarsia and Eretmocerus (Hymenoptera: Aphelinidae) are important biological control agents of whiteflies, and some of them not only parasitize hosts but also kill them with strong host‐feeding capacity. Two whitefly parasitoid species, Encarsia sophia and Eretmocerus melanoscutus were examined to determine if mating and host density affected their host feeding and parasitism. The whitefly host, Bemisia tabaci, was presented to these two wasp species in densities of 10, 20, 30, 40, 50 and 60 third‐instar nymphs per clip cage. Mated whitefly parasitoid females fed on more hosts than unmated females under a range of host densities (under all six host densities for En. sophia; under the densities of 40 nymphs or more for Er. melanoscutus). Meanwhile, mated females parasitized more whitefly nymphs than unmated females under all host densities for both species. With increase of host density, mated or unmated Er. melanoscutus females killed more hosts by host feeding and parasitism. Mated En. sophia females killed more hosts by host feeding with increase of host density, whereas unmated females did not parasitze whitefly nymphs at all. Our results suggest that only mated female parasitoids with host‐feeding behavior should be released in crop systems to increase their bio‐control efficiency.     

Parasites and supernormal manipulation.

Social parasites may exploit their hosts by mimicking other organisms that the hosts normally benefit from investing in or responding to in some other way. Some parasites exaggerate key characters of the organisms they mimic, possibly in order to increase the response from the hosts. The huge gape and extreme begging intensity of the parasitic common cuckoo chick (Cuculus canorus) may be an example. In this paper, the evolutionary stability of manipulating hosts through exaggerated signals is analysed using game theory. Our model indicates that a parasite's signal intensity must be below a certain threshold in order to ensure acceptance and that this threshold depends directly on the rate of parasitism. The only evolutionarily stable strategy (ESS) combination is when hosts accept all signallers and parasites signal at their optimal signal intensity, which must be below the threshold. Supernormal manipulation by parasites is only evolutionarily stable under sufficiently low rates of parasitism. If the conditions for the ESS combination are not satisfied, rejector hosts can invade using signal intensity as a cue for identifying parasites. These qualitative predictions are discussed with respect to empirical evidence from parasitic mimicry systems that have been suggested to involve supernormal signalling, including evicting avian brood parasites and insect-mimicking Ophrys orchids.     

The Population Dynamics and Community Ecology of Root Hemiparasitic Plants

Root hemiparasitic plants and their host plants interact directly, through parasitism, as well as indirectly, through scramble competition for resources. To understand the population dynamics and community ecology of root hemiparasitic plants and their hosts, models of resource-based competition have been extended to include resource parasitism. Parasitism provides a mechanism for parasitic plants to overcome deficits in their ability to compete for soil resources. The interaction ranges from competitive to exploiter-victim, depending on whether the benefits of parasitism overshadow the costs of competition. These models predict that as productivity in the system increases, parasitic plants should become more abundant. In diverse host communities, differences in the impact that parasites have on their hosts and the benefits that they receive from parasitizing different hosts may lead to nontransitive competitive relationships and a sort of apparent competition. The possible dynamics include paper-rock-scissors oscillations and indirect mutualisms between parasitic plants and their hosts that allow them to form coalitions that can exclude competitive dominants.     

Energetic cost of bot fly parasitism in free-ranging eastern chipmunks

The energy and nutrient demands of parasites on their hosts are frequently invoked as an explanation for negative impacts of parasitism on host survival and reproductive success. Although cuterebrid bot flies are among the physically largest and most-studied insect parasites of mammals, the only study conducted on metabolic consequences of bot fly parasitism revealed a surprisingly small effect of bot flies on host metabolism. Here we test the prediction that bot fly parasitism increases the resting metabolic rate (RMR) of free-ranging eastern chipmunks (Tamias striatus), particularly in juveniles who have not previously encountered parasites and have to allocate energy to growth. We found no effect of bot fly parasitism on adults. In juveniles, however, we found that RMR strongly increased with the number of bot fly larvae hosted. For a subset of 12 juveniles during a year where parasite prevalence was particularly high, we also compared the RMR before versus during the peak of bot fly prevalence, allowing each individual to act as its own control. Each bot fly larva resulted in a ~7.6% increase in the RMR of its host while reducing juvenile growth rates. Finally, bot fly parasitism at the juvenile stage was positively correlated with adult stage RMR, suggesting persistent effects of bot flies on RMR. This study is the first to show an important effect of bot fly parasitism on the metabolism and growth of a wild mammal. Our work highlights the importance of studying cost of parasitism over multiple years in natural settings, as negative effects on hosts are more likely to emerge in periods of high energetic demand (e.g. growing juveniles) and/or in harsh environmental conditions (e.g. low food availability).     

Long‐term coevolution between avian brood parasites and their hosts

Coevolutionary theory predicts that the most common long‐term outcome of the relationships between brood parasites and their hosts should be coevolutionary cycles based on a dynamic change selecting the currently least‐defended host species, given that when well‐defended hosts are abandoned, hosts will be selected to decrease their defences as these are usually assumed to be costly. This is assumed to be the case also in brood parasite‐host systems. Here I examine the frequency of the three potential long‐term outcomes of brood parasite–host coevolution (coevolutionary cycles, lack of rejection, and successful resistance) in 182 host species. The results of simple exploratory comparisons show that coevolutionary cycles are very scarce while the lack of rejection and successful resistance, which are considered evolutionary enigmas, are much more frequent. I discuss these results considering (i) the importance of different host defences at all stages of the breeding cycle, (ii) the role of phenotypic plasticity in long‐term coevolution, and (iii) the evolutionary history of host selection. I suggest that in purely antagonistic coevolutionary interactions, such as those involving brood parasites and their hosts, that although cycles will exist during an intermediate phase of the interactions, the arms race will end with the extinction of the host or with the host acquiring successful resistance. As evolutionary time passes, this resistance will force brood parasites to use previously less suitable host species. Furthermore, I present a model that represents the long‐term trajectories and outcomes of coevolutionary interactions between brood parasites and their hosts with respect to the evolution of egg‐rejection defence. This model suggests that as an increasing number of species acquire successful resistance, other unparasitized host species become more profitable and their parasitism rate and the costs imposed by brood parasitism at the population level will increase, selecting for the evolution of host defences. This means that although acceptance is adaptive when the parasitism rate and the costs of parasitism are very low, this cannot be considered to represent an evolutionary equilibrium, as conventional theory has done to date, because it is not stable.     

Insecticide mixtures for mosquito net impregnation against malaria vectors

Insecticides belonging to the pyrethroid family are the only compounds currently available for the treatment of mosquito nets. Unfortunately, some malaria vector species have developed resistance to pyrethroids and the lack of alternative chemical categories is a great concern. One strategy for resistance management would be to treat mosquito nets with a mixture associating two insecticides having different modes of action. This study presents the results obtained with insecticide mixtures containing several proportions of bifenthrin (a pyrethroid insecticide) and carbosulfan (a carbamate insecticide). The mixtures were sprayed on mosquito net samples and their efficacy were tested against a susceptible strain of Anopheles gambiae, the major malaria vector in Africa. A significant synergism was observed with a mixture containing 25 mg/m2 of bifenthrin (half the recommended dosage for treated nets) and 6.25 mg/m2 of carbosulfan (about 2% of the recommended dosage). The observed mortality was significantly more than expected in the absence of any interaction (80% vs 41%) and the knock-down effect was maintained, providing an effective barrier against susceptible mosquitoes.     

Frequency-dependent host selection by parasitic mites: a model and a case study

Previous studies on frequency-dependent food selection (changing food preferences in response to changes in relative food abundance) have focused on predators and parasitoids. These organisms utilize several victims during their lifetime. We introduce the case of parasites which, having accepted a host, do not change it. We propose two alternative models to explain the biased occurrence of parasites on different host types: (1) through the option of rejecting less-preferred hosts prior to accepting one of them; (2) by differential parasite survival on different host types. These models predict that host rejection, but not differential survival, can create frequency-dependent parasitism (FDP). Unlike previously described factors responsible for frequency dependence of food selection, which act through changing the foraging behaviour of individual predators or parasitoids, FDP involves no adjustment of parasite foraging strategy according to previous feeding experience. The mite Hemisarcoptes coccophagus is an obligate parasite of armoured scale insects (Homptera: Diaspididae). Our field data show that H. coccophagus is found more frequently on ovipositing than on young host females. Our model, combining the effects of host rejection and differential survival, is used to estimate the relative contribution of these factors to parasite biased occurrence on different hosts. The contribution of differential survival was dominant in H. coccophagus, and overode any effect of host rejection. Nevertheless, our prediction that FDP may be found in parasites is supported by literature data about a parasitic water mite.     

Relic behaviours, coevolution and the retention versus loss of host defences after episodes of avian brood parasitism

Most previous studies of brood parasitism have stressed that host defences, such as egg recognition, are lost in the absence of parasitism. Such losses could result in coevolutionary cycles in which parasites shift away from well-defended hosts only to switch back to them later at a time when these hosts have lost much or all of their defences and the parasite's current hosts have built up effective defences. However, the alternative 'single trajectory' model predicts that parasites rarely switch back to old hosts because ex-hosts retain egg recognition for long periods in the absence of parasitism. If true, egg recognition by the host may be a 'relic behaviour', because in the absence of parasitism its adaptive value is close to neutral. Using artificial nonmimetic eggs, I tested for egg recognition in two populations that are currently unparasitized but that are descended from lineages likely to have been parasitized in the past: the grey catbird, Dumetella carolinensis, on Bermuda and the loggerhead shrike, Lanius ludovicianus, in California. Both of these populations showed long-term retention, ejecting nonmimetic eggs at rates of nearly 100%. Because potential present-day selection pressures, such as conspecific parasitism, do not explain this egg recognition, Bermuda catbirds apparently retain recognition from North American conspecifics that were cowbird hosts before colonizing Bermuda and shrikes retain recognition from Old World congeners that were hosts of cuckoos. Retention is also indicated by passerines in California and the Caribbean that had high rejection rates of nonmimetic eggs before coming into contact with cowbirds. These new data suggest that both the coevolutionary cycles and single trajectory models have importance and that rejection behaviour can have insignificant costs, which is consistent with evolutionary lag explanations for the acceptance of parasitic eggs shown by some cuckoo and many cowbird hosts. Copyright 2001 The Association for the Study of Animal Behaviour.     

Density-dependent parasitoid recruitment per parasitized host: effects on parasitoid-host dynamics

Models of parasitoid-host dynamics are analyzed that include direct density dependence in the host population and either parasitoid- or host-density-dependent variation in parasitoid recruitment per parasitized host (parasitoid "yield"). The principal question addressed is how these forms of density dependence in parasitoid dynamics combine with aggregated parasitism to affect the stability of the models, in relation to suppression of host abundance. When parasitoid yield is an overcompensating function of either parasitoid or host density, stability is enhanced for systems with host equilibria suppressed far below the host carrying capacity. Substantially less aggregation of parasitism is required for stability in this situation than in previous models assuming parasitoid yield is constant. However, this density dependence in parasitoid yield also reduces stability when the host equilibrium is suppressed only moderately below carrying capacity; this is especially true when parasitoid yield is more strongly decreased by high host density than is host per capita reproduction. At present there is little empirical evidence concerning the relationships of parasitoid recruitment to parasitoid and host densities. The substantial effects shown in these models suggest that these relationships should be considered in empirical studies.     

Modeling the consequence of increased host tolerance toward avian brood parasitism

Avian brood parasites greatly reduce the reproductive success of their hosts. Empirical studies have demonstrated that some hosts have evolved defenses against parasitism like an ability to recognize and reject parasitic eggs that are dissimilar to their own eggs. Detailed mechanisms of how hosts recognize parasitism still remain unknown, but recent studies have shown that the host’s recognition, in many cases, is based on discordance of the eggs in a clutch, and that hosts are more error-prone when the nest is multiply parasitized, i.e., hosts tend to accept more multiple parasitism than single parasitism. In an area in Hungary, the great reed warbler Acrocephalus arundinaceus, one of the main hosts of the common cuckoo Cuculus canorus, is heavily parasitized and the parasitism rate has been kept at quite a high level for decades. Previous mathematical models suggest that such a high parasitism rate can be maintained because the focal host population behaves as a sink where few hosts can reproduce but immigration from outside replenishes the loss of host reproduction in the sink population. Here, we explore the consequences of the increased host tolerance towards multiple parasitism which has been overlooked in the previous studies using a simple model. Our model analysis shows that the increased host tolerance can dramatically contribute to both the parasite abundance and the parasitism rate being kept at a high level. We suggest that such a host behavior, combined with host immigration, can be an important factor responsible for the observed severe parasitism.     

Ectoparasites and age-dependent survival in a desert rodent

Host age is one of the key factors in host–parasite relationships as it possibly affects infestation levels, parasite-induced mortality of a host, and parasite distribution among host individuals. We tested two alternative hypotheses about infestation pattern and survival under parasitism in relation to host age. The first hypothesis assumes that parasites are recruited faster than they die and, thus, suggests that adult hosts will show higher infestation levels than juveniles because the former have more time to accumulate parasites. The second hypothesis assumes that parasites die faster than they are recruited and, thus, suggests that adults will show lower infestation levels because of acquired immune response and/or the mortality of heavily infested juveniles and, thus, selection for less infested adults. As the negative effects of parasites on host are often intensity-dependent, we expected that the age-related differences in infestation may be translated to lower or higher survival under parasitism of adults, in the cases of the first and the second hypotheses, respectively. We manipulated ectoparasite numbers using insecticide and assessed the infestation pattern in adult and juvenile gerbils (Gerbillus andersoni) in the Negev Desert. We found only a partial support for age-dependent parasitism. No age-related differences in infestation and distribution among host individuals were found after adjusting the ectoparasite numbers to the host’s surface area. However, age-related differences in survival under parasitism were revealed. The survival probability of parasitized juveniles decreased in about 48% compared to unparasitized hosts while the survival probability of adults was not affected by ectoparasites. Our results suggest that the effect of host age on host–parasite dynamics may not explicitly be determined by age-dependent differences in ectoparasite recruitment or mortality processes but may also be affected by other host-related and parasite-related traits.     

Method of insecticide delivery affects horizontal transfer of fipronil in the German cockroach (Dictyoptera: Blattellidae).

Horizontal transmission of insecticide occurs when foragers contact or ingest an insecticide, return to the aggregation or nest, and translocate the insecticide to the shelter and its vicinity. Relatively more sedentary members of the population then contact or eat the translocated insecticide and die. We evaluated three different methods of delivering fipronil to adult male German cockroaches, Blattella germanica (L.), for their potential to cause such secondary mortality in various developmental stages of the cockroach. Adult males topically treated with 5 ng of fipronil (approximately LD99) caused low mortality in untreated nymphs and no mortality in untreated adults within the same aggregation. Males exposed to residual fipronil on a glass surface translocated more insecticide, resulting in higher mortality of cockroaches they contacted, but only early instars were affected and no adult mortality was observed. Ingested fipronil bait, however, was most effectively translocated, and caused high mortality of untreated adults and nymphs. Ingestion of fipronil also caused greater secondary kill compared with a topical application of 25 ng, approximately the same amount recovered from the exterior of males that ingested 1 mg of 0.05% fipronil bait. Secondary mortality in the untreated population was significantly affected by the duration of contact between the treated and untreated cockroaches, the quantity and freshness of excretions from the treated insects, and the accessibility of the secretions to untreated cockroaches. The mechanisms that cause secondary kill may include ingestion of excreted fipronil residues, cannibalism of bait-fed cockroaches, as well as contact with fipronil-contaminated substrates.     

Population fluctuations of cabbage aphid,Brevicoryne brassicae (L.) (Hom.: Aphididae) and its natural parasitism rate on different canola cultivars

The cabbage aphid, Brevicoryne brassicae is one of the most destructive pests in Cruciferaceae or Brassicaceae plant family around the world. According to this pest rapid resistance to insecticide, using the resistant cultivars should be evaluated. In the present study, the different canola cultivars resistance against the cabbage aphid and the population fluctuations in canola fields and also the natural parasitism are evaluated in Shahre-rey region. To evaluate the population fluctuations and its natural parasitism on the RGS003, SLMOO46, Karag2, Licord, Sarigol, Opera, Ocapi and Talaei canola cultivars were cultivated in four replications using the complete randomised block design. The population fluctuations studies showed that the pest is active during the canola growth period in studied region, but the peak of population and damage is found in 6 May month. The results showed that natural parasitism percentage of cabbage aphid on different canola cultivars has not significant difference, but the parasitism amount in different sampling dates was significantly different, so that the natural parasitism maximum observed in date 29 April. These results showed that the different canola cultivars have significant effects on cabbage aphid and parasitoids activity amount. Using the present results, the recognised cultivars in integrated pest management programmes can be used to grow the laboratory aphids as the parasitoids hosts.     

Do distances among host patches and host density affect the distribution of a specialist parasitoid?

The effect of spatial habitat structure and patchiness may differ among species within a multi-trophic system. Theoretical models predict that species at higher trophic levels are more negatively affected by fragmentation than are their hosts or preys. The absence or presence of the higher trophic level, in turn, can affect the population dynamics of lower levels and even the stability of the trophic system as a whole. The present study examines different effects of spatial habitat structure with two field experiments, using as model system the parasitoid Cotesia popularis which is a specialist larval parasitoid of the herbivore Tyria jacobaeae. One experiment examines the colonisation rate of the parasitoid and the percentage parasitism at distances occurring on a natural scale; the other experiment examines the dispersal rate and the percentage parasitism in relation to the density of the herbivore and its host plant. C. popularis was able to reach artificial host populations at distances up to the largest distance created (at least 80 m from the nearest source population). Also, the percentage parasitism did not differ among the distances. The density experiment showed that the total number of herbivores parasitised was higher in patches with a high density of hosts, regardless of the density of the host plant. The percentage parasitism, however, was not related to the density of the host. The density of the host plant did have a (marginally) significant effect on the percentage parasitism, probably indicating that the parasitoid uses the host plant of the herbivore as a cue to find the herbivore itself. In conclusion, the parasitoid was not affected by the spatial habitat structure on spatial scales that are typical of local patches.